bobcat (lynx rufus · bobcat (lynx rufus) ecology in a longleaf pine ecosystem in southwestern...

BOBCAT (LYNX RUFUS) ECOLOGY IN A LONGLEAF PINE ECOSYSTEM IN

SOUTHWESTERN GEORGIA

by

JORDONA DOUGHTY

(Under the Direction of Robert J. Warren)

ABSTRACT

A paucity of knowledge exists about the effects of northern bobwhite (quail; Colinus

virginianus) management practices on bobcat (Lynx rufus) ecology. Quail management is an

important part of the longleaf pine (Pinus palustris)-wiregrass (Aristida stricta) ecosystem. This

study investigated bobcat home range, habitat use, and dietary patterns in a longleaf pine-

wiregrass ecosystem managed for quail in southwestern Georgia. Male home ranges were larger

than female home ranges, and home range sizes varied seasonally for females, but not for males.

Bobcats selected habitats to include in their home range, but did not select habitats within the

home range. Agriculture was the most preferred habitat type, and edge was an important

component of habitat. Diet varied seasonally during year 1 and year 2, but not during year 3.

Bobcats were not a major predator of quail. Their primary prey items were cotton rats

(Sigmodon hispidus) and other rodents. Land management practices such as prescribed burning

and maintenance of food plots probably contributed to high quality habitat with ample prey; thus

certain quail management practices may impact bobcat ecology.

INDEX WORDS: bobcat, diet, habitat use, home range, longleaf pine, Lynx rufus, scat, southwestern Georgia



by

JORDONA DOUGHTY

B.S., University of Delaware, 2002

A Thesis Submitted to the Graduate Faculty of The University of Georgia in Partial Fulfillment

of the Requirements for the Degree

MASTER OF SCIENCE

ATHENS, GEORGIA

2004

© 2004

Jordona Doughty

All Rights Reserved



by

JORDONA DOUGHTY

Major Professor: Robert J. Warren

Committee: L. Michael Conner Steven B. Castleberry Ronald L. Hendrick

Electronic Version Approved: Maureen Grasso Dean of the Graduate School The University of Georgia December 2004

iv

But the wildest of all the wild animals was the Cat.

He walked by himself, and all places were alike to him.

--Rudyard Kipling, Just So Stories

v

ACKNOWLEDGEMENTS

I would first like to extend my deepest gratitude to my co-major professors, Dr. Robert

Warren and Dr. Mike Conner. I am very lucky to have worked with such wonderful mentors.

Dr. Warren, your enthusiasm and sense of humor were always encouraging, and your advice

about my project, coursework, thesis-writing, and job-searching was well-taken. Dr. Conner,

without your statistical expertise, my data analysis would not have been possible. Your

instruction in trapping, editorial comments on every draft of every thesis chapter, assistance in

the job hunt, and everything else you helped me get through are all greatly appreciated. I would

also like to thank my other committee members, Dr. Steven Castleberry and Dr. Ron Hendrick,

for input on my thesis.

There are many people at Ichauway who helped make the bobcat project possible since

its inception almost 4 years ago. Thank you to Ivy Godbois for essentially setting up all of the

field components of the project. Although I arrived at Ichauway well into the project, many

thanks go out to Jerry Wade, Raymond Varnum, Brad Cross, and Brandon Rutledge for all of the

trapping they did early on and before I arrived. Thank you to Bobby Bass for setting up scat

lines and retrieving animals that meandered over to the Longleaf Plantation. I would also like to

thank Bobby for being a good friend and always putting a smile on my face. Thanks to Mark

Melvin for keeping us updated on bobcats he trapped off-site and for always inquiring about my

fitness progress. Thanks to Arthur Sheffield and the other guys at the shop for always keeping

Black Mud up and running, one headlight and all.

vi

Without the help of the Wildlife Lab at Ichauway, there would have been fewer bobcats

radio-tracked. Micah Perkins, Amanda Subalusky, and Brent Howze helped capture quite a few

animals before I arrived. I would like to give an extra thanks to Brent for helping me with

trapping, for always keeping on the lookout for scat, and for giving me a good laugh during

many conversations in the lab. Thanks to Allison Reid, who threw herself into trapping during

the short time she was here. Without her extensive recapture efforts, many bobcats would have

been lost to radio-collar failure.

I would not have survived fieldwork and getting through my project without the guidance

and friendship of Jessica Cochrane. Thanks JCC, for showing me all of the ropes of telemetry,

trapping, work-up of bobcats, etc. Thank you too, for keeping my mom at ease by

accompanying me on all of my night runs, well after you were done with telemetry duty (lol).

Without your kindness, I never would have legally driven again! Your friendship is one I will

never forget.

Thank you to many friends who always kept me encouraged. Marsha Ward and Justin

Davis were my personal cheerleaders, even though they were 4 hours away. I would especially

like to thank Marsha for being one of the best friends I could ever have, without whose

friendship I ever would have made it living in Georgia. Thanks to my roommates, Tara Muenz,

Allison Reid, and Sarah Cathey, for being good friends and always asking how things were

going. Tara and Allison never complained when I made coffee or breakfast at 2 or 3 a.m., trying

to be as quiet as possible and ending up making a huge racket. I would also like to thank all of

my hometown and college friends who spent long hours on the phone giving me encouragement

when I needed it most. Thank you to Ron, Angela, and Kim Kirby, my second family, for

providing support and love when I was miles away from my own family.

vii

Although they think I am crazy, my family has finally accepted that I am working

towards my dreams. Without the love and support of my parents, I never would have gone after

my goals or succeeded this far in my life. Thanks Mom, for being so incredibly overprotective

that I wanted to scream, but also for letting me become my own person just the same. Dad, I am

so thankful for your cheerful outlook and enthusiasm for everything I do. Thank you to my sister

Taylor, who is already following in my footsteps and who keeps me greatly entertained.

Someday, Taylor, you'll have your horse and we can ride together. I would really like to thank

Steven, my brother, whose friendship and respect mean the world to me. I hope you know that I

am as proud of you as you are of me. Thank you to all of my family for your encouragement,

even when you didn't quite understand what I am doing and why I do it. I will just keep showing

you the pretty pictures.

I would like to give a special thanks to my fiancé, Brian Kirby, for his admiration,

support, love, and understanding during my project. Thanks for staying up half the night on

Friday to do telemetry after a long drive from Rincon, and for getting up early on weekends to

run traps with us. You always put me high up on a pedestal, even when I didn't think I belonged

there. I can only hope I will be as patient during your Master's work as you have been with me.

I can't express to you how glad I am that we didn't give up on each other.

The Joseph W. Jones Ecological Research Center, the Robert Woodruff Foundation, and

the University of Georgia provided financial support for the project. Vehicles were provided by

the Georgia Department of Natural Resources. Thank you for Black Mud, she's a great truck!

Again, Thank you to everyone!

viii

TABLE OF CONTENTS

Page

ACKNOWLEDGEMENTS.............................................................................................................v

LIST OF TABLES...........................................................................................................................x

LIST OF FIGURES ....................................................................................................................... iv

CHAPTER

1 INTRODUCTION, STUDY AREA, JUSTIFICATION, AND THESIS FORMAT ....1

2 BOBCAT HOME RANGES IN A LONGLEAF PINE ECOSYSTEM IN

SOUTHWESTERN GEORGIA..............................................................................23

3 FACTORS AFFECTING BOBCAT HABITAT USE IN A LONGLEAF PINE

ECOSYSTEM IN SOUTHWESTERN GEORGIA................................................43

4 BOBCAT DIETS IN A LONGLEAF PINE ECOSYSTEM IN SOUTHWESTERN

GEORGIA ...............................................................................................................67

5 CONCLUSIONS AND MANAGEMENT IMPLICATIONS.....................................94

APPENDICES .............................................................................................................................104

A MANAGEMENT ZONES ON ICHAUWAY, BAKER COUNTY, GEORGIA, 2000-

2004 .......................................................................................................................104

B MORPHOLOGICAL DATA COLLECTED FOR 50 ADULT BOBCATS

CAPTURED AND RADIO-COLLARED BETWEEN DECEMBER 2000 AND

MAY 2004, ICHAUWAY, BAKER COUNTY, GEORGIA ...............................106

ix

C ANNUAL AND SEASONAL ADAPTIVE KERNEL (ADK) AND MINIMUM

CONVEX POLYGON (MCP) HOME RANGE SIZE ESTIMATES FOR

BOBCATS ON ICHAUWAY, BAKER COUNTY, GEORGIA, 2001-2004 ......109

D DESCRIPTION OF FIVE BOBCAT DENS LOCATED IN A LONGLEAF PINE

ECOSYSTEM, ICHAUWAY, BAKER COUNTY, GEORGIA, 2002-2004 ......116

E DESCRIPTION OF BOBCAT MORTALITIES IN SOUTHWESTERN GEORGIA,

2001-2004..............................................................................................................118

x

LIST OF TABLES

Page

Table 2.1: Studies documenting bobcat home ranges (km2) in the southeastern United States

(MMA=Modified Minimum Area; MCP=Minimum Convex Polygon; ADK=Adaptive

Kernel) ............................................................................................................................42

Table 3.1: Habitat type distance ratios for second-order selection using seasonal home ranges for

female bobcats at Ichauway, Baker County, Georgia, 2001-2004 .................................61

Table 3.2: Habitat rankings based on pair-wise comparisons between habitat type distance ratios

for second-order habitat selection of female bobcats monitored on Ichauway, Baker

County, Georgia, 2001-2004...........................................................................................62

Table 3.3: Habitat type distance ratios for second-order selection using seasonal home ranges for

male bobcats at Ichauway, Baker County, Georgia, 2001-2004.....................................63


for second-order habitat selection of male bobcats monitored on Ichauway, Baker

County, Georgia, 2001-2004...........................................................................................64

Table 3.5: Habitat type distance ratios for third-order selection using seasonal home ranges for

male and female bobcats combined at Ichauway, Baker County, Georgia, 2001-2004 .65


for third-order selection of all bobcats monitored on Ichauway, Baker County, Georgia,

2001-2004 .......................................................................................................................66

xi

Table 4.1: Studies documenting bobcat diet by percent occurrence in the southeastern United

States, using the gastrointestinal tract (GI; stomach or intestines) or scat for analysis ..93

iv

LIST OF FIGURES

Page

Figure 2.1: Seasonal home range sizes for male and female bobcats (F01=Fall 2001;

W02=Winter 2002; S02=Spring 2002; Su02=Summer 2002; F02=Fall 2002;

W03=Winter 2003; S03=Spring 2003; Su03=Summer 2003; F03=Fall 2003;

W04=Winter 2004; S04=Spring 2004) on Ichauway, Baker County, Georgia, 2001-

2004 ...............................................................................................................................40

Figure 4.1: Annual percent occurrence of prey items consumed by bobcats based on scat analysis

on Ichauway, Baker County, Georgia, 2001-2004........................................................83

Figure 4.2: Seasonal percent occurrence of prey items consumed by bobcats based on analysis of

135 scats on Ichauway, Baker County, Georgia, 2001-2002 ........................................85





Figure 4.5: Seasonal percent occurrence of prey items consumed by bobcats (S01=Summer

2001; F01=Fall 2001; W02=Winter 2002; S02=Spring 2002; Su02=Summer 2002;

F02=Fall 2002; W03=Winter 2003; S03=Spring 2003; Su03=Summer 2003; F03=Fall

2003; W04=Winter 2004; S04=Spring 2004) based on analysis of 413 scats on

Ichauway, Baker County, Georgia, 2001-2004.............................................................91

1

CHAPTER 1

INTRODUCTION, STUDY AREA, JUSTIFICATION, AND THESIS FORMAT

2

INTRODUCTION

The mesopredator-release hypothesis predicts that in the absence of apex predators, a

population explosion of medium-sized omnivores, hereafter referred to as mesopredators, will

occur (Rogers and Caro 1998). Apex predators are thought to suppress mesopredators, resulting

in ecosystem stabilization and greater species diversity. Therefore, the presence of apex

predators may increase prey species survival by controlling mesopredator populations

(Palomares et al. 1995, Rogers and Caro 1998, Courchamp et al. 1999, Crooks and Soule 1999,

Henke and Bryant 1999). Lower mesopredator population density results in improved nesting

success in songbirds and game birds through reduced nest predation (Sovada et al. 1995, Rogers

and Caro 1998, Courchamp et al. 1999, Crooks and Soule 1999).

Bobcats (Lynx rufus) are considered an apex predator in certain forested ecosystems of

the southeastern U.S. (Conner et al. 2000). However, control of bobcats and other mammalian

predators is often suggested as a way to increase game bird abundance. Although bobcats

historically have been considered a major predator of northern bobwhite (quail; Colinus

virginianus), few studies have specifically analyzed bobcat food habits in areas managed for

quail.

On 2 quail plantations in southern Alabama, quail were not an important part of bobcat

diets despite a high quail density (Miller and Speake 1978). In fact, the study suggested that

bobcat predation on cotton rats (Sigmodon hispidus) might be beneficial to quail populations by

decreasing nest predation and competition for food plants (Miller and Speake 1978).

3

In southwestern Georgia, quail remains were found in 1.9% of all bobcat scats collected over a 2-

year period (Cochrane 2003). However, a different study of mammalian predators in Georgia

found that quail remains in bobcat diets varied over a 2-year period, ranging from 0% - 12.5% on

1 of the areas studied (Schoch 2003).

A paucity of knowledge exists about the effects of quail management practices on bobcat

ecology, and whether bobcats have detrimental impacts on quail populations. Quail hunting is an

important cultural and economic tradition in the Southeast, and quail management is an

important part of the longleaf pine (Pinus palustris)-wiregrass (Aristida stricta) ecosystem

(Burger et al. 1999, Boring 2001). Thus, determining the impacts of bobcats on quail and the

impacts of quail management on bobcats are important for wildlife managers and researchers.

Home range

The bobcat is a solitary carnivore with variable home range sizes influenced by numerous

factors including geographic region, sex, age, season, habitat quality, prey availability and

abundance, and possibly experience (time-in-residence) (Fendley and Buie 1986, Anderson

1987, Rucker et al. 1989, Sandell 1989, Conner et al. 1999). Bobcats tend to have lower

densities and larger home ranges in the northern parts of their range, which have been related to

climatic/environmental and prey availability differences (Litvaitis et al. 1986, Anderson 1987,

Lovallo and Anderson 1996). Home ranges of bobcats in the Southeast vary from 1.1 km2 for

females and 2.6 km2 for males in Alabama (Miller and Speake 1979) to 24.5 km2 for females and

64.2 km2 for males in Arkansas (Rucker et al. 1989). Estimates of composite bobcat home range

sizes in southwestern Georgia over 2 years were 2.8 + 0.6 km2 for females and 6.0 + 0.8 km2 for

males (Cochrane 2003).

4

Perhaps the most important determinant of bobcat home range size is the association

between habitat quality and prey abundance and availability (Anderson 1987). An inverse

relationship exists between home range size and prey density (Mares et al. 1976, Buie et al.

1979, Knick 1990). According to the bobcat habitat suitability index model, the suitability of

habitat is determined by the ability of the habitat to support prey populations (Boyle and Fendley

1987). Habitats that are more suitable for abundant prey densities are more likely to be included

in a bobcat’s home range.

Generally, male home range sizes exceed those of females by 2-3 times, and may be as

much as 5 times larger (Hall and Newsom 1976, Buie et al. 1979, Kitchings and Story 1979,

Whitaker et al. 1987). Male home range size is affected by the size of female home ranges and

the number of mating opportunities, whereas female home ranges appear to be regulated by

diversity, abundance, stability and distribution of prey populations (Anderson 1987, Sandell

1989). Intrasexual overlap of home ranges, particularly between females, does not occur

frequently in bobcats, but male home ranges may overlap several female home ranges (Marshall

and Jenkins 1966, Hall and Newsom 1976, Buie et al. 1979, Miller and Speake 1979, McCord

and Cordoza 1982, Whitaker et al. 1987).

Size of home range also may be affected by season, though few studies of bobcats in the

Southeast have documented seasonal variation. Males have the largest home range during the

breeding season, and females have the smallest home range during parturition and kitten-rearing

(Anderson 1987). Seasonal fluctuation in home range sizes also may relate to seasonal

differences in prey availability. Home ranges were smallest during the summer in Arkansas,

probably because prey abundance is greatest during the warmest months (Rucker et al. 1989).

5

In southwestern Georgia, home ranges were largest for males during spring and summer

(Cochrane 2003). However, in South Carolina, bobcat home range size did not fluctuate

seasonally (Fendley and Buie 1986).

Only one study has investigated the influence of experience, or time-in-residence, of

bobcats on home range size. In a 5-year study of bobcats in Mississippi, changes in home range

size relative to time-in-residence were sex-dependent (Conner et al. 1999). Male bobcat home

range sizes were smaller in previous-year analysis than in subsequent years, whereas female

home ranges were larger in the previous year compared to subsequent years. Males may increase

their home range as they become well-established in their territories over time, probably to

increase breeding opportunities (Conner et al. 1999). As females establish themselves in an area

and develop hunting skills and familiarity with their home range over time, they will decrease

their home range size, optimize a smaller area, and save energy. Such relationships between sex,

time-in-residence, and home range size are poorly understood, creating a need for further

research.

Habitat Use

Habitat use patterns vary throughout the geographic range of the bobcat, but prey

abundance is the major determinant in habitat selection (Pollack 1951, Anderson 1987, Rucker et

al. 1989). Other factors that influence habitat use include resting site availability, denning site

availability, protection from environmental extremes, dense cover for hunting and escape, and

freedom from disturbance (Pollack 1951, Young 1958, Bailey 1974, Kitchings and Story 1984,

Anderson 1987, Boyle and Fendley 1987). Experience (time-in-residence), activity versus

inactivity, and diurnal versus nocturnal time periods are other potential influences on bobcat

habitat use (Conner et al. 1999).

6

Generally, bobcats prefer early to mid-successional habitats and areas with dense cover or rocky

outcroppings (Young 1958, Hall and Newsom 1976, Miller and Speake 1979, Knowles 1985,

Anderson 1987). Studies have found that bobcats also use human-modified areas, such as

abandoned logging roads, pipelines and old agricultural fields (Hall and Newsom 1976,

Kitchings and Story 1978, Miller and Speake 1979, Rucker et al. 1989).

Bobcat prey in the Southeast, especially the cotton rat and the eastern cottontail

(Sylvilagus floridanus), are most abundant in dense areas of early to mid-successional grass/forb-

shrub vegetation (Boyle and Fendley 1987). The shrub interspersion is necessary to provide

essential cover for prey species (Schnell 1968). Primary prey of bobcats also occurs in high

densities in broomsedge (Andropogon spp.)-vine habitat, characterized by herbaceous species

interspersed with shrubs and shrubby vines such as blackberry (Rubus spp.), Japanese

honeysuckle (Lonicera japonica), and trumpet vine (Bignonia radicans) (Golley et al. 1965).

Prescribed burning is a quail management practice conducive to improving habitat for bobcat

prey species by reducing the amount of woody understory, and promoting an herbaceous

understory.

In the Southeast, habitats selected by bobcats include pine plantations and agricultural

areas (Conner et al. 1992, Cochrane 2003), bottomland hardwoods (Heller and Fendley 1986,

Cochrane 2003), hardwoods associated with drainages in forested uplands (Zwank et al. 1985,

Cochrane 2003), and mid-successional stages with dense growth of saplings, vines and briars in

the bottomland hardwoods (Hall and Newsom 1976). These habitats have more abundant prey

than mature pine forests and other available habitats. Habitat use varies by season and is most

likely in response to seasonal shifts in prey availability and climatic variation (Rolley and Warde

1985). Sex-related differences in habitat use also occur in bobcats.

7

Females use higher quality habitat than males because they require more prey within smaller

home ranges, especially with changing energy demands during kitten-rearing (Rolley and Warde

1985). Den site availability also influences habitat use in females (Bailey 1974).

Experience (time-in-residence) may influence habitat use in bobcats, but only one study

has investigated this relationship. Home range habitat composition did not change relative to

time-in-residence for male or female bobcats in Mississippi, but it seemed to change for males

(Conner et al. 1999). Although differences in habitat use were not detected, more studies of this

potential relationship are necessary. Differences in active versus inactive animals and diurnal

versus nocturnal time periods also may influence habitat use in bobcats, but such influences have

never been investigated.

Dietary Patterns

Predators generally select prey within certain size limits to optimize ease in capture and

energy returns (Rosenzweig 1966, McCord and Cordoza 1982). Bobcats concentrate kills on

prey from 150-5,500 g, including squirrels (Sciurus spp.), pocket gophers (Geomys spp.,

Thomomys spp.), rabbits, large rodents and opossum (Didelphis virginiana)-sized mammals

(Rosenzweig 1966, McCord and Cordoza 1982, Boyle and Fendley 1987). They consume larger

prey, including deer (Odocoileus spp.), less frequently (McCord and Cordoza 1982, Story et al.

1982, Anderson and Lovallo 2003). Bobcats primarily consume rabbits and hares, particularly

cottontail rabbits (Sylvilagus spp.), throughout their range (Anderson 1987). Rodents comprise

the remaining bulk of the diet, though species vary by habitat. Some ground-dwelling avian (i.e.,

game and nongame) and reptilian (i.e., snakes) species also are consumed.

8

Bobcats may exhibit functional responses to prey by switching dietary composition from one

primary prey species to another as the abundance of different species changes (Baker et al.

2001). Bobcats also ingest grass, either accidentally or intentionally as a purgative (Miller and

Speake 1978, Buttrey 1979).

Variation in food habits occurs geographically. In the Southeast, cotton rats and eastern

cottontails compose most of the diet during all seasons, and cotton rats replace eastern cottontails

as the primary prey source when they are more abundant (Beasom and Moore 1977, Miller and

Speake 1978, Boyle and Fendley 1987, Cochrane 2003). White-tailed deer (Odocoileus

virginianus) become an important food source in some southern regions, especially during fall-

winter when carrion from the hunting season is available, and in late spring-summer, when fawns

are available (Buttrey 1979, Story et al. 1982). Deer become an important food source during

periods of low density of preferred prey (Beasom and Moore 1977). In mountainous and

highland regions of the South, squirrels, pine voles (Microtus pinetorum) and some bird species

become important food sources (Buttrey 1979, Kitchings and Story 1979).

Sex and age-related differences in food habits occur based on individual body size of

bobcats. In Arkansas, females consume more rats and mice than males (Fritts and Sealander

1978), and in New Hampshire, males consume more white-tailed deer and fewer cottontails than

females and juveniles (Litvaitis et al. 1984). Such differences in food habits appear to decrease

intraspecific competition within bobcat populations (Rosenzweig 1966, Fritts and Sealander

1978).

9

Reproductive/Den Ecology

Timing of the bobcat reproduction season, comprised of breeding, parturition, and

nursing young, varies according to geographic location, climate, photoperiod and prey

availability (McCord and Cordoza 1982). Breeding begins earlier and continues longer in

southern regions (McCord and Cordoza 1982). In Mississippi, the reproductive season occurs

between 1 January and 30 April (Jackson and Jacobson 1987). The post-parturition period, in

which the female must provide prey to her kittens, occurs between 1 May and 31 August, and

kittens remain with their mother through the fall season, until approximately 31 December

(Jackson and Jacobson 1987).

The location of bobcat den sites appears to be related to prey availability; females are

limited to hunting prey in close proximity to unprotected kittens (Bailey 1979). Den sites have

been discovered in hollow logs, rocky outcrops (Gashwiler et al. 1961, Cochrane 2003), at the

base of tree stumps in timber-harvested areas (Kitchings and Story 1984), and in thickets and

brush piles (Anderson 1987, Cochrane 2003). Den sites in human-made structures, such as

abandoned buildings, also have been observed (Bailey 1974). Auxiliary den sites, used by the

female bobcat and her kittens once they are old enough to travel with her, have been described in

rocky areas and abandoned holes of other species such as the woodchuck (Marmota monax)

(Bailey 1979, Kitchings and Story 1984). Prescribed burning may create thickets and hollow

stumps, thereby increasing the availability of den sites for bobcats (Young 1958, Kitchings and

Story 1984).

10

Longleaf Pine-Wiregrass Ecosystem

The longleaf pine-wiregrass ecosystem historically covered almost 36.5 million ha in the

southeastern United States. Today, this unique, fire-dependent ecosystem exists only in scattered

patches totaling less than 81,000 ha in the Coastal Plains of several southern states. Almost 30%

of remaining longleaf forest is located in Georgia (Holliday 2001). Many remaining tracts of

longleaf pine-wiregrass communities exist on private lands managed for northern bobwhite using

prescribed fire. Only about 3,600 ha of remaining longleaf forest is old-growth, and 1,024 ha of

old-growth is found in Georgia (Holliday 2001). Timber harvesting, development of naval stores

and production of turpentine, and fire suppression associated with land settlement contributed to

such a vast decline of the ecosystem (Engstrom et al. 2001).

Fire is critical to maintaining longleaf pine-wiregrass ecosystems because it reduces

competing hardwood species and woody understory, improves nutrient flow, encourages

regeneration in forest gaps, and induces a flowering response in wiregrass (Engstrom et al.

2001). Many species of plants and animals are considered threatened or endangered in the

longleaf pine-wiregrass ecosystem because they are found only in these declining fire-dependent

communities (Engstrom et al. 2001). The longleaf pine-wiregrass ecosystem has received much

attention in recent years due to its rapid disappearance. Restoration efforts at the public and

private level have been initiated throughout the Southeast. Managing for quail through

prescribed burning contributes to promoting longleaf pine restoration. Thus, research devoted to

quail management practices and their relationship to longleaf pine ecosystems has developed.

11

Quail Management Practices

Typical quail management activities, such as a 2-year rotation prescribed fire, planting

agricultural crops, maintaining food plots, and supplemental feeding, may prove beneficial to

bobcats. Prescribed fire increases and maintains a dense herbaceous understory and early

successional habitat, providing abundant resources and habitat for many small mammals that are

bobcat prey species (Golley et al. 1965, Miller and Speake 1979). Burning also may contribute

to more suitable denning habitat for bobcats by creating thickets and hollow stumps, which are

considered good denning sites (Young 1958, Kitchings and Story 1984).

Planting agricultural crops and maintaining quail food plots increases edge, also

providing greater prey availability for bobcats (Hall and Newsom 1976, Miller and Speake

1978). Food plots are usually agricultural grain patches established as borders between existing

agricultural fields and woods. Supplemental feeding, usually concentrated in feeders or scattered

in low vegetation or existing food plots, causes increased density and decreased home range size

in prey species, potentially attracting more predators (Landers and Mueller 1986, Boutin 1990).

In southwestern Georgia, bobcats were 10 times closer than expected to supplemental food than

expected (Godbois et al. 2004).

STUDY AREA

Ichauway is a privately owned 11,735-ha research facility located in Baker County,

Georgia, 16 km south of Newton, Georgia. It is located in the Dougherty Plain physiographic

province in the southeastern Gulf Coastal Plain (Boring 2001). Ichauway is characterized by flat

to gently rolling karst topography, with elevations ranging from 27 to 61 m. It has hot, humid

summers and short, mild, wet winters, with average daily temperatures ranging from 11.1°C

(winter) to 27.2°C (summer). Average annual precipitation is 132 cm per year (Boring 2001).

12

Longleaf pine woodlands and limesink wetlands are the dominant habitat types at

Ichauway. Other habitats include mixed pine-hardwood areas, food plots, agricultural fields,

slash pine (P. elliottii) flatwoods, riparian hardwood hammocks, oak sandhill barrens, natural

and old-field loblolly pine (P. taeda) stands, grassy and cypress-gum (Taxodium ascendens,

Nyssa biflora) limesink ponds, creek swamps, forested wetlands, riverine areas, shrub-scrub

upland, and human/cultural (i.e., resident quarters) areas (Boring 2001). The understory is

dominated by wiregrass and old-field grasses (e.g., Andropogon spp.), but >1,000 vascular plant

species occur on the site (Goebel et al. 1997, Drew et al. 1998). Approximately 24 km of the

Ichawaynochaway Creek flows through the study area, and the Flint River forms almost 22 km

of Ichauway’s eastern boundary (Boring 2001).

The site is divided into multiple-use and conservation zones interspersed throughout the

land area. Multiple-use zones comprise approximately 60% of Ichauway, and prescribed fire,

supplemental feeding, and maintenance of food plots are the primary management activities in

multiple-use zones. Conservation zones, comprising the remaining 40% of the land area, are

managed for longleaf pine restoration. Bobcats occur in both zones on Ichauway.

Much of Ichauway is managed for the longleaf pine-wiregrass ecosystem with prescribed

fire. Burning is performed on a 2-year rotation, usually during winter and early spring, on

approximately 4,000 to 6,000 ha throughout the entire site (Godbois et al. 2004).

Prescribed burning is 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).

13

Food plots consisting of brown top millet (Brachiaria ramose), winter wheat (Triticum

aestivum), cowpea (Vigna spp.), grain sorghum (Sorghum vulgare), and Egyptian wheat

(Sorghum spp.) comprise 20% of the property (Godbois et al. 2004). Food plots occur less

abundantly in conservation zones than multiple use zones, and are typically planted for white-

tailed deer rather than quail. Supplemental feeding for quail with grain sorghum occurs in

multiple use zones at 2-week intervals between November and May (Godbois et al. 2004).

Fields are disked to improve quail food availability by allowing ragweed (Ambrosia

artemisiifolia) and partridge pea (Chamaecrista fasciculata) seedlings and other plants to grow

(Landers and Mueller 1986, Davis 2001).

Limited predator removal occurs in multiple use zones after the quail-hunting season

(March-May) annually. The primary predators removed are raccoons (Procyon lotor) and

opossums. Low numbers of coyote (Canis latrans), red fox (Vulpes vulpes), gray fox (Urocyon

cinereoargentus), and striped skunk (Mephitis mephitis) also are removed each year. Bobcats

were harvested occasionally before 1999, but since then have not been harvested.

JUSTIFICATION

Quail hunting makes a significant economic impact and it is part of a long-held cultural

tradition in the southeastern United States (Burger et al. 1999). Thus, quail management in the

South is important, and is a component of maintaining fire-dependent, longleaf pine-wiregrass

ecosystems. Because bobcats historically have been considered detrimental to game birds such

as quail through predation on adults and eggs, they often are harvested on quail plantations.

Research documenting the relationship between bobcats and quail is lacking but necessary for

quail plantation managers. Bobcats may be beneficial to quail by reducing populations of quail

predators or competitors.

14

Thus, a study of bobcat ecology in a quail management area is important for both researchers and

managers. Studying the ecology of bobcats in the longleaf pine-wiregrass ecosystem will aid in

future management decisions regarding bobcats in the southeastern U.S. If evidence suggests

that bobcats may have a stabilizing effect in longleaf pine-wiregrass ecosystems by regulating

other predators that may be detrimental to quail, then bobcat conservation may improve

restoration efforts for this threatened ecosystem.

Economic demand for non-endangered bobcat and lynx (Lynx canadensis) in North

America increased in the 1970s after the import of fur of endangered cats was made illegal

through passage of the Endangered Species Conservation Act of 1969 (Anderson 1987).

Between 1970 and 1976, annual harvest of bobcats across the United States increased 3-fold, and

price per pelt rose more than 10 times (Anderson 1987). Bobcats were added to Appendix II of

the Convention on International Trade in Endangered Species (CITES) in 1975 in response to a

rising concern about possible over-exploitation (Anderson 1987, Conner et al. 1992). As a result

of the CITES listing, bobcat research has become important because each state must monitor

populations to ensure that harvest does not prove detrimental to the species (Anderson 1987,

Conner et al. 1992). Because bobcat home range size, density, and habitat use vary

geographically, it is necessary to gather information regarding basic ecology on a regional basis

to make effective management decisions (Rucker et al. 1989, Conner et al. 1992). The ecology

of bobcats in Georgia has only recently been studied.

OBJECTIVES

1. Determine if bobcat home range size varies annually, and by season and sex.

2. Determine bobcat habitat use at 2 spatial scales, and compare bobcat habitat use

between sexes and among seasons.

15

3. Determine whether activity status and time-of-day influence bobcat habitat use.

4. Quantify bobcat diet and determine if it varies seasonally and annually.

THESIS FORMAT

The thesis was written in manuscript format. Chapters 2, 3, and 4 represent manuscripts

to be submitted for publication. Chapter 1 is an introduction to the thesis and summarizes prior

research of bobcat ecology. Chapter 2 describes bobcat home ranges in the longleaf pine-

wiregrass ecosystem and will be submitted to the American Midland Naturalist. Chapter 3

describes bobcat habitat use and will be submitted to the Journal of Wildlife Management.

Chapter 4 describes bobcat diet patterns and will be submitted to the Southeastern Naturalist.

Chapter 5 summarizes all findings and conclusions.

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Southeastern Association of Fish and Wildlife Agencies 46:147-158.

_____, B. W. Plowman, B. D. Leopold, and C. Lovell. 1999. Influence of time-in-residence on

home range and habitat use of bobcats. Journal of Wildlife Management 63:261-269.

_____, B. D. Leopold, and M. J. Chamberlain. 2000. Multivariate habitat models for bobcats in

southern forested landscapes. Pages 51-55 in Proceedings of a symposium on current

bobcat research and implications for management. The Wildlife Society 2000

Conference, Nashville, Tennessee, USA.

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mesopredator release effect. Journal of Animal Ecology 68:282-292.

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fragmented system. Nature 400:563-566

Davis, M. S. 2001. Creature feature: Northern bobwhite quail. Pages 19-29 in J. R. Wilson, ed.

The fire forest: longleaf pine/wiregrass ecosystem. Georgia Wildlife Federation Natural

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Drew, M. B., L. K. Kirkman, and A. K. Gholson, Jr. 1998. The vascular flora of Ichauway,

Baker County, Georgia: a remnant longleaf pine-wiregrass ecosystem. Castanea 63:1-

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Engstrom, T., L. K. Kirkman, and R. J. Mitchell. 2001. The natural history of the fire forest.

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Covington, GA.

Fendley, T. T. and D. E. Buie. 1986. Seasonal home range and movement patterns of the bobcat

on the Savannah River Plant. Pages 237-259 in S. D. Miller and D. D. Everett, eds. Cats

of the world: biology, conservation, and management. National Wildlife Federation,

Washington, D. C.

Fritts, S. H. and J. A. Sealander. 1978. Diets of bobcats in Arkansas with special reference to

age and sex differences. Journal of Wildlife Management 42:533-539.

Gashwiler, J. S., W. L. Robinette, and O. W. Morris. 1961. Breeding habits of bobcats in Utah.

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518.

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USA.

Golley, F. B., J. B. Gentry, L. D. Caldwell, and L. B. Davenport. 1965. Number and variety of

small mammals on the AEC Savannah River Plant. Journal of Mammalogy 46:1-18.

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Hall, H. T. and J. D. Newsom. 1976. Summer home ranges and movements of bobcats in

bottomland hardwoods of southern Louisiana. Proceedings of the Annual Conference

Southeastern Association of Fish and Wildlife Agencies 30: 427-436.

Heller, S. P. and T. T. Fendley. 1986. Bobcat habitat on the Savannah River Plant, South

Carolina. Pages 415-423 in S. D. Miller and D. D. Everett, eds. Cats of the world:

biology, conservation, and management. National Wildlife Federation, Washington,

D.C.

Henke, S. E. and F. C. Bryant. 1999. Effects of coyote removal on the faunal community in

western Texas. Journal of Wildlife Management 63:1066-1081.

Holliday, P. P. 2001. Going, going…saving the longleaf pine ecosystem before it’s gone. Pages

55-66 in J. R. Wilson, ed. The fire forest: longleaf pine-wiregrass ecosystem.

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Covington, GA.

Jackson, D. L. and H. A. Jacobson. 1987. Population ecology of the bobcat (Felis rufus) in

managed southern forest ecosystems. Final Rep. Fed. Aid. Proj. W-48-30, 31, 32, 33, 32.

Miss. Dep. Wildl. Conserv. Study XX. 69 pp.

Kitchings, J. T. and J. D. Story. 1978. Preliminary studies of bobcat activity patterns.

Proceedings of the Annual Conference Southeastern Association of Fish and Wildlife

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_____ and _____. 1979. Home range and diets of adult bobcats in eastern Tennessee. Pages 47-


conference. National Wildlife Federation Scientific and Technical Series 6.

20

_____ and _____. 1984. Movements and dispersal of bobcats in eastern Tennessee. Journal of

Wildlife Management 48:957-961.

Knick, S. T. 1990. Ecology of bobcats relative to exploitation and a prey decline in southeastern

Idaho. Wildlife Monographs 108. 42 pp.

Knowles, P. R. 1985. Home range size and habitat selection of bobcats, Lynx rufus, in north-

central Montana. Canadian Field Naturalist 99:6-12.

Landers, J. L. and B. S. Mueller. 1986. Bobwhite quail management: a habitat approach. Tall

Timbers Research Station and Quail Unlimited.

Litvaitis, J. A., C. L. Stevens, and W. W. Mautz. 1984. Age, sex, and weight of bobcats in

relation to winter diets. Journal of Wildlife Management 48:632-635.

_____, J. A. Sherburne, and J. A. Bissonette. 1986. Bobcat habitat use and home range size in

relation to prey density. Journal of Wildlife Management 50:110-117.

Lovallo, M. J. and E. M. Anderson. 1996. Bobcat (Lynx rufus) home range size and habitat use

in northwest Wisconsin. American Midland Naturalist 135:241-252.

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striatus. Oecologia 25:1-12.

Marshall, A. D. and J. H. Jenkins. 1966. Movements and home ranges of bobcats as determined

by radio-tracking in the upper coastal plain of South Carolina. Proceedings of the Annual

Conference of Southeastern Association of the Game and Fish Commission 20:206-214.

McCord, C. M. and J. E. Cordoza. 1982. Bobcat and lynx. Pages 728-766 in J. A. Chapman

and G. A. Feldhamer, eds. Wild Mammals of North America. Johns Hopkins University

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21

Miller, S. D. and D. W. Speake. 1978. Prey utilization on quail plantations in southern

Alabama. Proceedings of the Annual Conference of the Southeastern Association of Fish

and Wildlife Agencies 32:100-111.

_____ and _____. 1979. Progress report: Demography and home range of the bobcat in south

Alabama. Pages 123-124 in L. G. Blum and P. C. Escherich, eds. Proceedings of the

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top predators by controlling smaller predator populations: an example with lynx,

mongoose, and rabbits. Conservation Biology 9:295-305.

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the mesopredator release hypothesis. Oecologia 116:227-233.

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Rucker, R. A., M. L. Kennedy, G. A. Heidt, and M. J. Harvey. 1989. Population density,

movements, and habitat use of bobcats in Arkansas. Southwestern Naturalist 34:101-108.

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182 in J. L. Gittleman, ed. Carnivore behavior, ecology and evolution. Cornell

University Press, Ithaca, New York, USA.

22

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populations. Journal of Wildlife Management 32:698-711.

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red foxes on duck nest success. Journal of Wildlife Management 59:1-9.

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Kentucky bobcats. Proceedings of the Annual Proceedings of the Southeastern

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23

CHAPTER 2

BOBCAT HOME RANGES IN A

LONGLEAF PINE ECOSYSTEM IN SOUTHWESTERN GEORGIA¹

_______________________ ¹Doughty, J., J. C. Cochrane, I. A. Godbois, L.M. Conner, and R. J. Warren. 2004. To be submitted to the American Midland Naturalist.

24

ABSTRACT.─Little is known about bobcat (Lynx rufus) ecology in areas managed for northern

bobwhite (quail; Colinus virginianus). Therefore, we determined seasonal and annual home

range sizes of bobcats in a longleaf pine (Pinus palustris) forest managed for quail in

southwestern Georgia. We monitored 44 radio-collared bobcats during 2001-2004. Average

annual home range size was 11.0 + 1.4 km2 for male bobcats and 6.4 + 1.0 km2 for females.

There was no sex × year interaction (F1,25 = 0.15, P = 0.700). Annual home ranges differed

between sexes (F1,25= 7.54, P = 0.011) but not among years (F1,25 = 2.79, P = 0.107). Male

bobcats had a mean home range size of 8.5 + 1.0 km2 across seasons. The smallest male home

range (2.1 + 3.3 km2) occurred during fall 2001, and the largest home range (15.9 + 2.7 km2)

occurred during winter 2002. Females had a mean home range size of 5.3 + 0.7 km2 across

seasons. The smallest female home range (2.8 + 1.7 km2) occurred during summer 2002 and the

largest home range (8.5 + 1.7 km2) occurred during winter 2003. For seasonal home ranges,

there was a significant sex × season interaction (F10,181 = 1.64, P = 0.100); thus, we examined

seasonal home ranges for the sexes separately. Home range sizes varied seasonally for females

(F10,124 = 3.22, P = 0.001), but not for males (F10,52 = 0.88, P = 0.554). Home ranges of both

sexes were smaller than home ranges previously reported for the southeastern United States.

Land management practices such as prescribed burning and maintenance of food plots

contributed to high quality habitat with ample prey, which probably led to smaller home range

sizes of bobcats on this study area compared to other studies.

25

INTRODUCTION

The bobcat (Lynx rufus) is a solitary carnivore with variable home range sizes influenced

by numerous factors including geographic region, sex, age, season, habitat quality, prey

availability and abundance, and possibly experience (time-in-residence) (Fendley and Buie,

1986; Anderson, 1987; Rucker et al., 1989; Sandell, 1989; Conner et al., 1999). Home ranges of

bobcats in the Southeast vary from 1.1 km2 for females and 2.6 km2 for males (Miller and

Speake, 1979) to 24.5 km2 for females and 64.2 km2 for males (Rucker et al., 1989). Estimates

of composite bobcat home range sizes in southwestern Georgia over 2 years were 2.8 + 0.6 km2

for females and 6.0 + 0.8 km2 for males (Cochrane, 2003).

Male home ranges typically exceed those of females by 2-3 times, and may be as much as

5 times larger (Hall and Newsom, 1976; Buie et al., 1979; Kitchings and Story, 1979; Whitaker

et al., 1987). Male home range size is affected by the size of female home ranges and the

number of mating opportunities, whereas female home ranges appear to be regulated by

diversity, abundance, stability, and distribution of prey populations (Anderson, 1987; Sandell,

1989). Male home ranges may overlap several female and male home ranges, but intrasexual

overlap between females appears rare (Marshall and Jenkins, 1966; Hall and Newsom, 1976;

Buie et al., 1979; Miller and Speake, 1979; McCord and Cordoza, 1982; Whitaker et al., 1987).

However, some studies reported frequent overlap among female home ranges (Zezulak and

Schwab, 1979; Chamberlain and Leopold, 2001; Nielsen and Woolf, 2001).

Differences in habitat quality are the most often used explanation for home range

variability in bobcats (Anderson, 1987). According to the bobcat habitat suitability index model,

the suitability of habitat is determined by the ability of the habitat to support prey populations

(Boyle and Fendley, 1987).

26

Studies have documented an inverse relationship between prey abundance and home range size

(Buie et al., 1979, Knick, 1990). Habitats more suitable for abundant prey densities are more

likely to be included in a bobcat’s home range, and higher quality habitat should result in smaller

home ranges (Buie et al., 1979, Knick, 1990). Conner et al. (2001) suggested that habitat quality

influenced bobcat home range size, but once habitat quality increases to a threshold, home ranges

become influenced by other factors such as density and breeding opportunities.

Home range size also may vary seasonally, though few studies of bobcats in the

Southeast have documented seasonal variation. Males have the largest home range during the

breeding season, and females have the smallest home range during parturition and kitten-rearing

(Anderson, 1987; Knick, 1990; Conner et al., 1992). Seasonal fluctuation in home range sizes

also may relate to seasonal differences in prey availability. Home ranges were smallest during

the summer in Arkansas, probably because prey abundance is greatest during the warmest

months (Rucker et al. 1989). In southwestern Georgia, home ranges were largest for males

during spring and summer (Cochrane, 2003). However, in South Carolina, bobcat home range

size did not fluctuate seasonally (Fendley and Buie, 1986).

Although seasonal home ranges of bobcats in the longleaf pine ecosystem have been

studied, annual home ranges have not been estimated. Our objectives were to determine annual

and seasonal home range sizes of bobcats in a longleaf pine ecosystem in southwestern Georgia.

We hypothesized that male home ranges would be larger than female home ranges, and that

seasonal variation would occur.

27

STUDY AREA






















28



approximately 4,000 to 6,000 ha throughout the entire site (Godbois et al. 2004). Prescribed

burning is 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).












opossums (Didelphis virginiana). Low numbers of coyote (Canis latrans), red fox (Vulpes

vulpes), gray fox (Urocyon cinereoargentus), and striped skunk (Mephitis mephitis) also are

removed each year. Bobcats were harvested occasionally before 1999, but since then have not

been harvested.

29

METHODS

Bobcat capture, handling, and monitoring.─We trapped bobcats with baited #3 Victor Soft

Catch traps (Woodstream Corp., Lititz, PA), and baited #1.75 Oneida Victor coil-spring traps

(Victor Inc., Ltd., Cleveland, OH). Animals were captured from December 2000 until May

2004, though trapping efforts were sporadic between July 2001 and October 2003.

Captured animals were netted and given an intramuscular injection of ketamine hydrochloride

(10 mg/kg body weight) (Seal and Kreeger, 1987). We recorded sex, weight, total body length,

hind foot length, ear length, and tail length, and classified animals as adult or juvenile based on

secondary sex characteristics, length, and weight (Crowe, 1975). Adults were fitted with a 180-g

VHF radio-collar (Advanced Telemetry Systems, Isanti, MN). Each bobcat received a uniquely

numbered ear tattoo. Beginning in November 2003, 3-mm ear punches were taken from every

captured animal for a concurrent genetic study. Bobcats were monitored and released 8 to 24

hours after sedation at the trap site to ensure full recovery. All trapping procedures were

approved by the University of Georgia Institutional Animal Care and Use Committee (IACUC

#A990159).

Using radio telemetry, we began monitoring bobcats 2-7 days after release. We obtained

locations by triangulation, taking ≥2 radio telemetry azimuth locations from known reference

points with a 3-element Yagi antenna (Sirtrack, New Zealand) and hand-held receiver (Wildlife

Materials Inc., Carbondale, IL). To minimize error due to animal movement between readings,

time between consecutive bearings was ≤15 minutes (Cochran, 1980; Kenward, 1987; White and

Garrott, 1990).

30

Each bobcat was located 4-6 times per week, and locations were obtained equally throughout the

diel period, with ≥8 hours between each location to ensure biological independence. We

determined activity (active or inactive) by a change in the pulse rate of the transmitter or signal

intensity when movement was detected (Chamberlain et al., 1998).

Data Analysis.─We used the FORTRAN program EPOLY (L. M. Conner, pers. comm.) to

convert radio telemetry locations into Universal Transverse Mercator (UTM) coordinates. We

calculated 95% adaptive kernel (ADK; Worton, 1989) annual and seasonal home ranges for

bobcats with ≥30 locations per calendar season (annual = 12 consecutive months; fall = 21 Sep-

20 Dec; winter = 21 Dec-20 Mar; spring = 21 Mar-20 Jun; summer = 21 Jun-20 Sep) using

CALHOME (Kie et al., 1996). We also calculated minimum convex polygon (MCP) home

range estimates for comparison with other studies (Mohr, 1947). Annual home ranges were

determined for animals monitored for 4 consecutive seasons.

Statistical analyses were only performed on ADK home range estimates. To determine

whether annual home range differed as a function of sex, year, or a sex × year interaction, we

used an Analysis of Variance (ANOVA) with PROC GLM (SAS Institute, 2003). We used a

repeated measures ANOVA with PROC MIXED to determine whether seasonal home range size

differed as a function of sex, season, or the interaction (SAS Institute, 2003). Animals were

treated as the subject, repeated over seasons. We considered statistical significance at α=0.10.

RESULTS

We radio-tracked 13 - 27 bobcats each season from 21 September 2001-20 June 2004 (44

total animals, 17M and 27F; 29 annual home ranges). Male bobcat (11.0 + 1.4 km2) ADK

annual home ranges were almost 2 times larger (F1,25 = 7.54, P = 0.011) than female (6.4 + 1.0

km2) annual home ranges.

31

Annual home ranges did not differ (F1,25 = 2.79, P = 0.107) among years. There was no sex ×

year interaction (F1,25 = 0.15, P = 0.700). According to MCP estimates, males (8.2 + 1.2 km2)

had a mean annual home range approximately 1.5 times larger than females (5.2 + 1.1 km2).

For seasonal home ranges, there was a significant sex × season interaction (F10,181 = 1.64,

P = 0.100); thus, we examined seasonal home ranges for the sexes separately. Home range sizes

varied seasonally for females (F10,124 = 3.22, P = 0.001), but not for males (F10,51.6 = 0.88, P =

0.554) (Figure 2.1). Male bobcats had a mean home range size of 8.5 + 1.0 km2 across seasons,

and females had a mean home range size of 5.3 + 0.7 km2 across seasons. For males, the

smallest home range (2.1 + 3.3 km2) occurred during fall 2001, and the largest home range (15.9

+ 2.7 km2) occurred during winter 2002. The smallest female home range (2.8 + 1.7 km2)

occurred during summer 2002 and the largest home range (8.5 + 1.7 km2) occurred during winter

2003.

DISCUSSION

Similar to most studies of bobcat home ranges in the southeastern U.S., we found that

male bobcats had larger home ranges than females (Table 2.1). However, home ranges of both

sexes were smaller than home ranges previously reported, probably the result of difference in

habitat quality among study areas (Kitchings and Story, 1979; Buie et al., 1979; Hamilton, 1982;

Shiftlet, 1984; Lancia et al., 1986; Rucker et al., 1989; Conner et al., 1992; Conner et al., 2001).

The ability of a habitat to support prey populations determines the suitability of bobcat habitat

(Boyle and Fendley, 1987). High prey abundance should result in smaller home ranges, because

bobcats in habitats that maintain high prey densities do not have to travel as far to fulfill their

dietary needs (Buie et al., 1979; Knick, 1990).

32

Prescribed fire was a primary land management tool on our study site, which increases

and maintains a dense herbaceous understory and early successional habitat, ultimately providing

abundant resources and habitat for prey populations (Golley et al. 1965; Miller and Speake,

1979). Between 4,000 and 6,000 ha are burned annually on the study area, providing ample

habitat with early-successional herbaceous vegetation. Approximately 20% of the study area is

made up of wildlife food plots and agriculture (Godbois et al., 2004). Planting agricultural crops

and maintaining quail food plots increases edge, which also provides ample resources for prey

(Hall and Newsom, 1976; Miller and Speake, 1978; Cummings and Vessey, 1994). In addition,

approximately 270 metric tons of grain sorghum are spread over 7,020 ha throughout areas on

Ichauway that are managed for quail between November and May each year (Godbois et al.,

2004). In a preliminary analysis of small mammal data collected on our study area, cotton rats

(Sigmodon hispidus) were 5.5 times greater, house mouse (Mus musculus) were 3.5 times

greater, cotton mouse (Peromyscus gossypinus) were 1.5 times greater, and Eastern harvest

mouse (Reithrodontomys humulis) were 2 times greater in supplementally-fed versus unfed areas

(L. M. Conner, Joseph W. Jones Ecological Research Center, unpublished data). Thus, current

land management practices on our study site may have influenced bobcat home range sizes by

concentrating prey populations and concomitantly establishing high quality bobcat habitat.

Seasonal home range sizes differed according to sex. Male home ranges were larger than

females during all seasons except fall 2001. An exceptionally large home range of 1 female

(Bobcat #27, 18.4 km2) may have contributed to the average female home range for fall 2001

being larger than the average male home range size during that season. Although male home

range sizes did not vary seasonally, we observed that males had the largest home ranges during

winter in all 3 years.

33

Similar to other studies, this finding suggests that male bobcats increased their home ranges

during the breeding period, which occurs during the winter months in the southeastern states

(Anderson, 1987; Jackson and Jacobson, 1987; Knick, 1990; Conner et al. 1992). Increasing

their home range during the reproductive season allows males more breeding opportunities by

overlapping more female home ranges (Anderson and Lovallo, 2003).

Female home range sizes varied seasonally. The average home range size during winter

2003 was significantly larger than 9 of the other 10 seasons, which likely explains the seasonal

variation in home range size for female bobcats on our study site. It is possible that during the

colder months prey were more difficult to find, and expansion of the home range was necessary.

However, the home range of several females during that season probably caused the difference.

Bobcat #'s 40, 6, and 18 had home range sizes of 34.7 km2, 17.5 and 14.0 km2, respectively,

which were considerably larger than other females.

Between winter and spring during all 3 years of the study, female home range sizes

declined, which was likely due to females restricting their movements during denning (Bailey,

1974; Knick, 1990). The smallest female home range occurred during summer 2002, which

corresponded to the post-parturition period during that year, in which the female must provide

prey to her kittens (Bailey, 1979; Jackson and Jacobson, 1987; Conner et al., 1992). Home

ranges increased in size between summer and fall seasons, during which kittens become old

enough to travel with their mother (Bailey, 1979).

We concluded that home ranges of bobcats in southwestern Georgia were smaller than

most home ranges previously reported for the Southeast. Land management practices such as

prescribed burning, supplemental feeding, and maintenance of food plots likely contributed to

high quality habitat with ample prey.

34

High quality habitat containing abundant prey resources probably contributed to smaller home

range sizes of bobcats in the longleaf pine ecosystem compared to most other studies. Future

research should document prey population densities and bobcat locations in burned versus non-

burned areas and in food plots versus non-food plots, which might provide stronger evidence of

the role land management practices play in bobcat ecology in the longleaf pine forest.

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patterns of adult bobcats in Central Mississippi. Proc. Annu. Conf. Southeast. Assoc. of

Fish and Wildl. Agencies, 52:191-196.

_____ and B. D. Leopold. 2001. Spatio-Temporal relationships among adult bobcats in central

Mississippi. Pages 45-50 In: Woolf, A., C. K. Nielsen, and R. D. Bluett, (eds.). Proc.

Symposium on Current Bobcat Res. and Impl. for Manage., The Wildlife Society 2000


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_____, B. W. Plowman, B. D. Leopold, and C. Lovell. 1999. Influence of time-in-residence on

home range and habitat use of bobcats. J. Wildl. Manage., 63:261-269.

_____, M. J. Chamberlain, and B. D. Leopold. 2001. Bobcat home range size relative to habitat

quality. Proc. Annu. Conf. Southeast. Assoc. of Fish and Wildl. Agencies, 55:418-426.

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Crowe, D. M. 1975. Aspects of aging, growth, and reproduction of bobcats from Wyoming.

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Davis, M. S. 2001. Creature feature: Northern bobwhite quail. Pages 19-29 In: J. R. Wilson,

(ed.). The fire forest: longleaf pine-wiregrass ecosystem. Georgia Wildlife Federation

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Fendley, T. T. and D. E. Buie. 1986. Seasonal home range and movement patterns of the bobcat

on the Savannah River Plant. Pages 237-259 In: S. D. Miller and D. D. Everett, (eds.).

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Golley, F. B., J. B. Gentry, L. D. Caldwell, and L. B. Davenport. 1965. Number and variety

of small mammals on the AEC Savannah River Plant. J. Mammal., 46:1-18.

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Island, South Carolina. Thesis, University of Georgia, Athens, Georgia, USA. 84 pp.

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bottomland hardwoods of southern Louisiana. Proc. Annu. Conf. Southeast. Assoc. of

Fish and Wildl. Agencies, 30: 427-436.

Hamilton, D. A. 1982. Ecology of the bobcat in Missouri. Thesis, University of Missouri,

Columbia, Missouri, USA. 152 pp.


managed southern forest ecosystems. Final Rep. Fed. Aid. Proj., W-48-30, 31, 32,

33, 32. Miss. Dep. Wildl. Conserv. Study XX. 69 pp.

Kenward, R. 1987. Wildlife radio tagging. Harcourt Brace Jovanovich. London, England. 222

pp.

Kie, J. G., J. A. Baldwin, and C. J. Evans. 1996. CALHOME: A program for estimating animal

home range. Wildl. Soc. Bull., 24:342-344.

Kitchings, J. T. and J. D. Story. 1979. Home range and diets of adult bobcats in eastern

Tennessee. Pages 47-52 In: L. G. Blum and P.C. Escherich, (eds.). Proc. Bobcat Res.

Conf., National Wildlife Federation Scientific and Technical Series 6.


Idaho. Wildl. Mono., 108. 42 pp.

Lancia, R. A., D. K. Woodward, and S. D. Miller. 1986. Summer movement patterns and

habitat use by bobcats on Croatan National Forest, North Carolina. Pages 425-436 In:

S. D. Miller and D. D. Everett, (eds.). Cats of the world: biology, conservation, and

management. Caesar Kleberg Wildlife Research Institute, Kingsville, Texas, USA.


Timbers Research Station and Quail Unlimited, Tallahassee, Florida, USA.

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by radio-tracking in the upper coastal plain of South Carolina. Proc. Annu. Conf.

Southeast. Assoc. of Game and Fish Comm., 20:206-214.

McCord, C. M. and J. E. Cordoza. 1982. Bobcat and lynx. Pages 728-766 In: J. A. Chapman

and G. A. Feldhamer, (eds.). Wild mammals of North America. Johns Hopkins

University Press, Baltimore, Maryland, USA.


Alabama. Proc. Annu. Conf. Southeast. Assoc. of Game and Fish Comm., 32:100-111.

_____ and _____. 1979. Progress report: demography and home range of the bobcat in south

Alabama. Pages 123-124 In: L. G. Blum and P.C. Escherich, (eds.). Proc. Bobcat Res.


Nielsen, C. K. and A. Woolf. 2001. Spatial organization of bobcats (Lynx rufus) in southern

Illinois. Am. Midl. Nat., 146:43-52.

Mohr, C. O. 1947. Table of equivalent populations of North American small mammals. Am.

Midl. Nat., 37:223-249.


movements, and habitat use of bobcats in Arkansas. Southwest. Nat., 34:101-108.

Sandell, M. 1989. The mating tactics and spacing patterns of solitary carnivores. Pages 164-

182 In: J. L. Gittleman, (ed.). Carnivore behavior, ecology and evolution. Cornell

University Press, Ithaca, New York, USA.

SAS Institute, Inc. 2003. SAS User's Guide: Statistics, 2003 edition. SAS Inst. Inc., Cary,

North Carolina, USA.

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Seal, U. S., and T. J. Kreeger. 1987. Chemical immobilization of furbearers. Pages 191–215 In:

M. Novak, J. A. Baker, M. E. Obbard, and B. Malloch, (eds.). Wild furbearer

management and conservation in North America. Ministry of Natural Resources,

Ontario, Canada.

Shiftlet, B. L. 1984. Movements, activity, and habitat use of the bobcat in upland mixed pine-

hardwoods. Thesis, Louisiana State University, Baton Rouge, Louisiana, USA.

Whitaker, J., R. B. Fredrick, and T. L. Edwards. 1987. Home-range size and overlap of eastern

Kentucky bobcats. Proc. Annu. Conf. Southeast. Assoc. of Fish and Wildl. Agencies,

41:417-423.

White, G. C. and R. A. Garrott. 1990. Analysis of wildlife radio-tracking data. Academic Press,

Inc., San Diego, California. 383 pp.

Worton, B. J. 1989. Kernel methods for estimating the utilization in home-range studies.

Ecol., 70:164-168.

Zezulak, D. S. and R. G. Schwab. 1979. A comparison of density, home-range, and habitat

utilization of bobcat populations at Lava Beds and Joshua Tree National Monuments,

California. Pages 74-79 In: L. G. Blum and P.C. Escherich, (eds.). Proc. Bobcat Res.


40

Figure 2.1. Seasonal home range sizes for male and female bobcats (F01=Fall 2001;

W02=Winter 2002; S02=Spring 2002; Su02=Summer 2002; F02=Fall 2002; W03=Winter 2003;

S03=Spring 2003; Su03=Summer 2003; F03=Fall 2003; W04=Winter 2004; S04=Spring 2004)

on Ichauway, Baker County, Georgia, 2001-2004.

41

seasons

F01 W02 S02 Su02 F02 W03 S03 Su03 F03 W04 S04

hom

e ra

nge

size

(km

2 )

-2

0

2

4

6

8

10

12

14

16

18

20

22

Males

Females

42

Table 2.1. Studies documenting bobcat home range sizes (km²) in the southeastern United States (MMA=Modified Minimum Area; MCP=Minimum Convex Polygon; ADK=Adaptive Kernel). ______________________________________________________________________________ Reference State Sample Home range Home range size M F model ______________________________________________________________________________

Hall and Newsom, 1976a LA 6 4.9 1.0 MMA

Kitchings and Story, 1979 TN 5 42.9 11.5 MCP

Miller and Speake, 1979 AL 20 2.6 1.1 MCP

Buie et al., 1979b SC 6 20.8 10.3 MCP

Hamilton, 1982 MO 30 60.4 16.1 MCP

Shiftlet, 1984 MS 7 10.1 5.9 MCP

Fendley and Buie, 1986 SC 7 3.2 1.6 MCP

Lancia et al., 1986d NC 8 37.7 22.1 MCP

Rucker et al., 1989 AR 6 64.2 24.5 MCP

Conner et al., 1992 MS 15 36.5 20.6 MCP

Griffin, 2001 SC 8 10.5-16.7 3.5-10.5 ADK

Conner et al., 2001 MS 42 20.2 12.3 MCP

This study GA 29 8.2 5.2 MCP This study GA 29 11.0 6.4 ADK ______________________________________________________________________________ a Used only summer data. b Used only fall and winter data.

43

CHAPTER 3

FACTORS AFFECTING BOBCAT HABITAT USE

IN A LONGLEAF PINE ECOSYSTEM IN SOUTHWESTERN GEORGIA¹

_______________________ ¹Doughty, J., J. C. Cochrane, I. A. Godbois, L.M. Conner, and R. J. Warren. 2004. To be submitted to the Journal of Wildlife Management.

44

Abstract: Little is known about bobcat (Lynx rufus) ecology in areas managed for northern

bobwhite (quail; Colinus virginianus). We investigated bobcat habitat use in a longleaf pine

(Pinus palustris) forest managed for quail in southwestern Georgia. We monitored 43 radio-

collared bobcats from 2001 – 2004 to determine habitat use at 2 spatial scales: second order

(habitat selection of home range within the study site) and third order (habitat selection within an

individual’s home range). We also investigated whether activity status (active or inactive) of

individuals or time-of-day (day, night, or crepuscular) affected habitat use. At the second order

of selection, there was no sex × season interaction (Λ = 0.804, P = 1.000), and season had no

effect (Λ = 0.708, P = 0.783). Male and female bobcats selected habitat differently (Λ = 0.919,

P = 0.033). Female bobcats were closer to all habitat types than expected, suggesting a

preference for edges. Females preferred agriculture over all other habitat types. Male bobcats

were closer to agriculture, hardwood, pine regeneration, pine, mixed hardwood, and urban/barren

habitat types than expected, and they preferred agriculture over all other habitat types. At the

third order of selection, there was no sex × season interaction (Λ = 0.786, P = 0.998). Bobcat

habitat selection did not differ by sex (Λ = 0.947, P = 0.210) or season (Λ = 0.654, P = 0.306).

When data for sex and season were pooled, overall habitat selection occurred (Λ = 0.596,

P = 0.012). Urban/barren was most preferred, followed by wetland, hardwood, agriculture,

shrub/scrub, pine regeneration, pine, and mixed pine-hardwood. Bobcats did not select habitat

differently according to activity status (Λ = 0.990, P = 0.981) or time-of-day (Λ = 0.972,

P = 0.647). We suggest that bobcats on our study site prefer agricultural areas and other habitats

that provide a dense herbaceous layer because they produce abundant prey. Prescribed fire,

interspersed food plots, and other quail management practices create habitats preferred by

bobcats.

45

Key Words: activity status, bobcat, Georgia, habitat use, Johnson's second order selection,

Johnson's third order selection, longleaf pine, Lynx rufus, time-of-day

______________________________________________________________________________

INTRODUCTION

Habitat use patterns vary throughout the geographic range of the bobcat (Lynx rufus), but

prey abundance is the major determinant in habitat selection (Pollack 1951, Anderson 1987,

Rucker et al. 1989). Other factors that influence habitat use include resting site availability,

denning site availability, protection from environmental extremes, dense cover for hunting and

escape, and freedom from disturbance (Pollack 1951, Young 1958, Bailey 1974, Kitchings and

Story 1984, Anderson 1987, Boyle and Fendley 1987). Activity status and time-of-day are

potential influences on bobcat habitat use, but such influences have not been studied.

Shrub interspersion is necessary to provide cover for prey species (Schnell 1968).

Bobcat prey in the Southeast, especially the cotton rat (Sigmodon hispidus) and the eastern

cottontail (Sylvilagus floridanus), are most abundant in dense areas of early to mid-successional

grass/forb-shrub vegetation (Boyle and Fendley 1987). Cotton rats and other rodents also occur

in high densities in broomsedge (Andropogon spp.)-vine habitat, characterized by herbaceous

species interspersed with shrubs and shrubby vines such as blackberry (Rubus spp.), Japanese

honeysuckle (Lonicera japonica), and trumpet vine (Bignonia radicans) (Golley et al. 1965), and

they are commonly found near agriculture and food plots (Cummings and Vessey 1994). In the

Southeast, habitats selected by bobcats include pine plantations and agricultural areas (Conner et

al. 1992, Cochrane 2003), and bottomland hardwoods (Heller and Fendley 1986, Cochrane

2003).

46

Bobcats also prefer hardwoods associated with drainages in forested upland (Zwank et al. 1985),

and mid-successional stages with dense growth of saplings, vines and briars in bottomland

hardwoods (Hall and Newsom 1976). These habitats have more abundant prey than mature pine

forests and other available habitats.

There is a paucity of information about bobcat habitat use in a longleaf pine (Pinus

palustris)-wiregrass (Aristida stricta) ecosystem. Therefore, the objectives of this study were to

determine bobcat habitat use at Johnson's second and third orders of selection (Johnson 1980),

and to determine whether activity status and time-of-day affect habitat use. We hypothesized

that bobcats would select early successional habitat types and edges, that active animals would

be located more than expected in early successional habitat types, and that early successional

habitats would be used more than expected during night and crepuscular periods.

STUDY AREA









slash pine (P. elliottii) flatwoods, riparian hardwood hammocks, and oak sandhill barrens.

47

Natural and old-field loblolly pine (P. taeda) stands, grassy and cypress-gum (Taxodium

ascendens, Nyssa biflora) limesink ponds, creek swamps, forested wetlands, riverine areas,

shrub-scrub upland, and human/cultural (i.e., resident quarters) areas are also found at Ichauway

(Boring 2001). The understory is dominated by wiregrass and old-field grasses (e.g.,

Andropogon spp.), but >1,000 vascular plant species occur on the site (Goebel et al. 1997, Drew

et al. 1998). Approximately 24 km of the Ichawaynochaway Creek flows through the study area,

and the Flint River forms almost 22 km of Ichauway’s eastern boundary (Boring 2001).
















tailed deer rather than quail.

48

Supplemental feeding for quail with grain sorghum occurs in multiple use zones at 2-week

intervals between November and May (Godbois et al. 2004). Fields are disked to improve quail

food availability by allowing ragweed (Ambrosia artemisiifolia) and partridge pea

(Chamaecrista fasciculata) seedlings and other plants to grow (Landers and Mueller 1986, Davis

2001).






been harvested.

METHODS

Bobcat capture, handling, and monitoring

We trapped bobcats with baited #3 Victor Soft Catch traps (Woodstream Corp., Lititz,

PA), and baited #1.75 Oneida Victor coil-spring traps (Victor Inc., Ltd., Cleveland, OH).

Animals were captured from December 2000 until May 2004, though trapping efforts were

sporadic between July 2001 and October 2003. Captured animals were netted and given an

intramuscular injection of ketamine hydrochloride (10 mg/kg body weight) (Seal and Kreeger

1987). We recorded sex, weight, total body length, hind foot length, ear length, and tail length,

and classified animals as adult or juvenile based on secondary sex characteristics, length, and

weight (Crowe 1975).

We fitted adults with a 180-g VHF radio-collar (Advanced Telemetry Systems, Isanti,

MN). Each bobcat received a uniquely numbered ear tattoo.

49

Beginning in November 2003, 3-mm ear punches were taken from every captured animal for a

concurrent DNA study. We monitored and released bobcats 8 to 24 hours after sedation at the

trap site to ensure full recovery from sedation. All trapping procedures were approved by the

University of Georgia Institutional Animal Care and Use Committee (IACUC #A990159).

Using radio telemetry, we began monitoring bobcats 2-7 days after release. We obtained

locations by triangulation, taking ≥2 radio telemetry azimuth locations from known reference

points with a 3-element Yagi antenna (Sirtrack, New Zealand) and hand-held receiver (Wildlife

Materials Inc., Carbondale, IL). Each bobcat was located 4-6 times per week, and locations were

obtained equally throughout the diel period, with ≥8 hours between each location to ensure

biological independence. To minimize error due to animal movement between readings, time

between consecutive bearings was ≤15 minutes (Cochran 1980, Kenward 1987, White and

Garrot 1990). We determined activity (active or inactive) by a change in the pulse rate of the

transmitter or signal intensity when movement was detected (Chamberlain et al. 1998).

Data Analysis

We used the FORTRAN program EPOLY (L. M. Conner, Joseph W. Jones Ecological

Research Center, unpublished data) to convert radio telemetry locations into Universal

Transverse Mercator (UTM) coordinates. We calculated 95% minimum convex polygon (MCP;

Mohr 1947) annual and seasonal home ranges for bobcats with ≥30 locations per calendar season

(e.g., fall = Sep-Dec; winter = Dec-Mar; spring = Mar-Jun; summer = Jun-Sep) using

CALHOME (Kie et al. 1996). We performed all habitat analyses using ARC/INFO and

Geographic Information System (GIS) software (ESRI 2004).

50

Habitat was classified into 8 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 from aerial photo-interpretation ARC/INFO into

a GIS. Bobcat locations and home ranges were intersected onto habitat maps using ARC/INFO.

We determined habitat use at 2 spatial scales: Johnson's second order (habitat selection of

home range within the study site) and Johnson's third order (habitat selection within an

individual’s home range) (Johnson 1980). We used a Euclidean distance technique to test for

habitat selection by comparing the mean distance between animal locations and habitat types to

corresponding expected distances (Conner and Plowman 2001, Conner et al. 2003).

Random locations for the study area and for each home range were generated, and we

calculated the mean distance (m) from random locations in the study area to each habitat type

and random locations within each home range to each habitat type using the NEAR command in

ARC/INFO (ESRI 2004). We also calculated mean distances from each bobcat location to each

habitat type in the home range. For second order selection, we created 8 distance ratios for each

bobcat (1 for each habitat type): the average distances from random locations within the home

range to each habitat type divided by the average distances from random locations in the study

area to each habitat type. For third order selection, we created 8 additional distance ratios—the

average distances from bobcat locations within the home range (i.e., used distances) to each

habitat type divided by the average distances from random locations within the home range (i.e.,

random distances) to each habitat type.

51

We assessed differences in habitat use as a function of sex, season, and their interaction

using a 2-factor multivariate analysis of variance (MANOVA). 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, the habitat type was preferred; if

the ratio was >1, the habitat type was avoided. We used univariate t-tests on each habitat type to

determine disproportional habitat use if the distance ratio did not equal 1. Ranking matrices

were created using univariate t-tests (i.e., pairwise mean comparisons) to rank habitat types.

All statistical analyses were performed with SAS software (SAS Institute, 2003). We considered

statistical significance at α=0.10.

We also investigated whether activity status and time-of-day affected habitat selection

seasonally. We defined an active animal as one that was observed moving (by a change in the

transmitter motion switch or signal intensity) during one or both radio-telemetry bearings for

each location. We defined daytime as the time from 2 hours after sunrise to 2 hours before

sunset on the median day of each calendar season, and nighttime as the time from 2 hours after

sunset to 2 hours before sunrise on the median day of each calendar season, taking into account

Daylight Saving Time during the appropriate seasons. We defined crepuscular periods as the

time period 2 hours before and 2 hours after sunrise and sunset. Seasons were pooled across

years (e.g., Fall = fall 2001, 2002 and 2003). We analyzed differences in habitat use according

to activity, sex and season using the Euclidean distance technique and a 3-factor MANOVA.

Similarly, we used a 3-factor MANOVA to detect differences in habitat use according to time-

of-day, sex, and season.

52

RESULTS

We radio-tracked 13 - 27 bobcats each season from 21 September 2001-20 June 2004 (43

total animals, 16M and 27F). At the second order of selection, there was no sex × season

interaction (Λ = 0.804, P = 1.000), and season had no effect (Λ = 0.708, P = 0.783). Therefore,

seasonal data were combined to analyze the sexes separately. Male and female bobcats selected

habitat differently (Λ = 0.919, P = 0.033). Female bobcats were closer to all habitat types than

expected (Table 3.1), suggesting a preference for edges. Females preferred agriculture over all

other habitat types, followed by mixed pine-hardwood, pine regeneration, hardwood, wetland,

shrub/scrub, urban/barren, and pine (Table 3.2). Male bobcats were closer to agriculture,

hardwood, pine regeneration, pine, mixed pine-hardwood, and urban/barren habitat types than

expected (Table 3.3). They preferred agriculture over all other habitat types, followed by mixed

pine-hardwood, pine, hardwood, urban/barren, wetland, pine regeneration, and shrub/scrub

(Table 3.4). Males significantly preferred agriculture over shrub/scrub, pine regeneration,

wetland, and urban/barren.

At the third order of selection, there was no sex × season interaction (Λ = 0.786, P =

0.998). Bobcats habitat selection did not differ by sex (Λ = 0.947, P = 0.210) or season

(Λ = 0.654, P = 0.306). When data for sex and season were pooled, overall habitat selection

occurred (Λ = 0.596, P = 0.012). Bobcats were farther than expected from pine regeneration,

pine, and mixed pine-hardwood, but were closer than expected to wetland and urban/barren

habitat (Table 3.5). Urban/barren was most preferred, followed by wetland, hardwood,

agriculture, shrub/scrub, pine regeneration, pine, and mixed pine-hardwood (Table 3.6).

Contrary to our hypotheses, bobcats did not select habitat differently according to activity status

(Λ = 0.990, P = 0.981) or time-of-day (Λ = 0.972, P = 0.647).

53

DISCUSSION

In the longleaf pine ecosystem of Ichauway, bobcats selected habitats to include in their

home range (Johnson's second order of selection), but did not select habitats within the home

range (Johnson's third order of selection). At Johnson's second order, female bobcats were closer

to all habitat types (agriculture, shrub/scrub, hardwood, pine regeneration, pine, mixed pine-

hardwood, wetland and urban/barren) than expected, and males were closer then expected to all

habitat types except shrub/scrub and wetland. According to Conner et al. (2003), animals prefer

edge when closer to all habitat types than expected. Thus, bobcats appeared to prefer edge when

establishing home ranges. Edge provides travel routes and access to hunting areas such as

agriculture fields, and it is typically where an abundance of prey is found (Landers and Mueller

1986, Cummings and Vessey 1994).

Similar to other studies in the Southeast, our bobcats preferred agriculture (Cochrane

2003, Conner et al. 1992), mixed pine-hardwood (Cochrane 2003), pine (Conner et al. 1992), and

hardwood habitats (Hall and Newsom 1976, Lancia et al. 1986). Bobcats likely selected early

and mid-successional habitats due to prey abundance and availability. Cotton rats and eastern

cottontails, two primary bobcat prey species in the Southeast, are typically most abundant in

dense areas of early to mid-successional grass/forb-shrub vegetation (Boyle and Fendley 1987).

Agricultural areas (e.g. food plots and edges) also attract rodent and lagomorph species

(Cummings and Vessey 1994). Hardwood and mixed pine-hardwood habitats contain a dense

herbaceous understory and shrub interspersion necessary to provide essential cover for prey

species, and such habitats also provide bobcats with cover (Golley et al. 1965, Schnell 1968).

54

On our study area, the mixed pine hardwood and hardwood habitat types are commonly

associated with wet areas near the creek and river. Thus, in addition to providing suitable

resources for bobcats and their prey, these habitats also provided cool, shady areas for refuge

(Godbois 2003).

We observed sex-related differences in habitat use at Johnson's second order. Females

preferred agriculture over all other habitat types, whereas males preferred agriculture over

shrub/scrub, pine regeneration, wetland, and urban/barren. Females preferred pine regeneration

more than males, whereas males preferred pine more than females. Females typically use higher

quality habitat than males because they require more prey within smaller home ranges, especially

with increased energy demands during kitten-rearing (Rolley and Warde 1985). Den site

availability also influences habitat use in females, and potential den sites may be found in

hardwood and mixed pine-hardwood habitat types (Bailey 1974).

At Johnson's third order, bobcat habitat selection did not differ by sex, season, or their

interaction. Bobcats used agriculture, shrub/scrub, hardwood, and wetland habitat types as

expected. They were farther than expected from pine regeneration, pine, and mixed pine-

hardwood. However, bobcats were closer than expected to urban/barren habitat, probably

because 1 female and 1 male (Bobcat #10 and #34, respectively) maintained home ranges in the

vicinity of the laboratory, research, and residential buildings on Ichauway, where most of the

urban/barren habitat was located. Habitat selection within the home range was likely not as

critical because high-quality habitat was selected when establishing the home range (Cochrane

2003).

55

Although bobcats are considered nocturnal, evidence suggests that they are most active

during the hours around sunrise and sunset, which corresponds with activity peaks of rodents and

lagomorphs, the primary prey species (Marshall and Jenkins 1966, Hall and Newsom 1976,

Kitchings and Story 1978, Buie et al. 1979, Anderson 1987, Chamberlain et al. 1998). We

expected active animals to select agriculture and other early-to-mid-successional habitat types

more than inactive animals. We also expected bobcats to prefer early-to-mid-successional

habitats during the nighttime and crepuscular periods. Contrary to our hypotheses, bobcats did

not select habitat differently according to activity status or time-of-day. Because bobcats

selected early-to-mid-successional habitat types to include in their home ranges and such habitats

may provide dense herbaceous cover, resting animals (i.e., inactive) likely did not have to find

cover in other habitat types. If preferred habitats provided ample prey and resting sites, then

bobcats probably did not have to move to other habitats during periods of rest or at different

times throughout the diel period.

MANAGEMENT IMPLICATIONS

Our findings suggest that bobcat habitat selection is probably determined by prey

availability. Rodents are commonly found in agricultural areas containing food plots, and

bobcats preferred agricultural habitats over all other habitat types. Habitat should be managed to

provide a dense herbaceous layer for prey and bobcats. Prescribed fire is the primary

management tool in the longleaf pine ecosystem that maintains such vegetation. In addition,

managing for a mosaic of diverse habitat types increases the amount of edge available, which is

an important component of bobcat habitat. Although we did not find activity status or time-of-

day as factors influencing habitat selection by bobcats, consideration of such factors may be

important to other species that are strictly nocturnal, diurnal, or crepuscular.

56

LITERATURE CITED










Reports 82(10.147).





Chamberlain, M. J., L. M. Conner, B. D. Leopold, and K. J. Sullivan. 1998. Diel activity

patterns of adult bobcats in Central Mississippi. Proceedings of the Annual Conference

of the Southeastern Association of Fish and Wildlife Agencies 52:191-196.

Cochran, W. W. 1980. Wildlife Telemetry. Pages 507-520 in S. D. Schemnitz, ed. Wildlife

management techniques manual. 4th ed. The Wildlife Society Inc., Washington, D. C.



57




_____ and B. W. Plowman. 2001. Using Euclidean distances to assess nonrandom habitat use.

Pages 275-290 in J. J. Millspaugh and J. M. Marzluff, eds. Radio tracking and animal

populations. Academic Press, San Diego, California, USA.

_____, M. D. Smith, and L. W. Burger. 2003. A comparison of distance-based and

classification-based analyses of habitat use. Ecology 84:526-531.

Crowe, D. M. 1975. Aspects of aging, growth, and reproduction of bobcats from Wyoming.


Cummings, J. R. and S. H. Vessey. 1994. Agricultural influences of movement patterns of

white-footed mice (Peromyscus leucopus). American Midland Naturalist 132:209-218.


The fire forest: longleaf pine-wiregrass ecosystem. Georgia Wildlife Federation Natural

Georgia Series 8(2). Georgia Wildlife Press, Covington, Georgia.


Baker County, Georgia: a remnant longleaf pine/wiregrass ecosystem. Castanea 63:1-

24.

Environmental Systems Research Institute (ESRI). 2004. ARC/INFO version 9.0.

Environmental Systems Research Institute, Redlands, California, USA.

Godbois, I. A. 2003. Ecology of bobcats on land managed for northern bobwhite in

southwestern Georgia. Thesis, University of Georgia, Athens, Georgia, USA.

58

_____, L. M. Conner, and R. J. Warren. 2004. Space-use patterns of bobcats relative to


518.


Technical Report 97-1. Joseph W. Jones Ecological Research Center, Newton, Georgia,

USA.




bottomland hardwoods of southern Louisiana. Proceedings of the Annual Conference of

the Southeastern Association of Fish and Wildlife Agencies 30: 427-436.

Heller, S. P. and T. T. Fendley. 1986. Bobcat habitat on the Savannah River Plant, South

Carolina. Pages 415-423 in S. D. Miller and D. D. Everett, eds. Cats of the world:

biology, conservation, and management. National Wildlife Federation, Washington, D.C.

Johnson, D. H. 1980. The comparison of usage and availability measurements for evaluating

resource preference. Ecology 61:65-71.

Kenward, R. 1987. Wildlife radio tagging. Harcourt Brace Jovanovich. London, England. 222

pp.

Kie, J. G., J. A. Baldwin, and C. J. Evans. 1996. CALHOME: A program for estimating animal

home range. Wildlife Society Bulletin 24:342-344.

Kitchings, J. T. and J. D. Story. 1978. Preliminary studies of bobcat activity patterns.

Proceedings of the Annual Conference of the Southeastern Association of Fish and

Wildlife Agencies 32:53-59.

59




habitat use by bobcats on Croatan National Forest, North Carolina. Pages 425-436 in S.

D. Miller and D. D. Everett, eds. Cats of the world: biology, conservation, and

management. National Wildlife Federation, Washington, D.C.




by radio-tracking in the upper coastal plain of South Carolina. Proceedings of the Annual

Conference of the Southeastern Association of Game and Fish Commissioners 20:206-

214.

Mohr, C. O. 1947. Table of equivalent populations of North American small mammals.

American Midland Naturalist 37:223-249.

Pollack, E. M. 1951. Observation on New England bobcats. Journal of Mammalogy 32:356-

358.

Rolley, R. E. and W. D. Warde. 1985. Bobcat habitat use in southeastern Oklahoma. Journal of






60

Seal, U. S., and T. J. Kreeger. 1987. Chemical immobilization of furbearers. Pages 191–215 in

M. Novak, J. A. Baker, M. E. Obbard, and B. Malloch, editors. Wild furbearer

management and conservation in North America. Ministry of Natural Resources,

Ontario, Canada.



White, G. C. and R. A. Garrott. 1990. Analysis of wildlife radio-tracking data. Academic Press,

Inc., San Diego, California. 383 pp.


193 pp.

Zwank, P. J., B. L. Shiflet, and J. D. Newsom. 1985. Habitat use by bobcats in upland forests

of Louisiana. Proceedings of the Annual Conference of the Southeastern Association of


61

Table 3.1. Mean habitat type distance ratios for second-order selection using seasonal home ranges for female bobcats at Ichauway, Baker County, Georgia, 2001-2004. _________________________________________________________________________ Habitat type Meana t Pb

_________________________________________________________________________ Agriculture -0.3473 -7.08 <0.0001

Shrub/Scrub -0.0700 -0.80 0.4301

Hardwood -0.1104 -1.82 0.0808

Pine regeneration -0.1301 -1.39 0.1753

Pine -0.0434 -0.44 0.6634

Mixed pine-hardwood -0.1616 -2.48 0.0198

Wetland -0.0091 -0.11 0.9140

Urban/Barren -0.0314 -0.42 0.6777 _________________________________________________________________________

aAverage distance from random locations within home ranges divided by average distance from random locations throughout study area. Mean ratios <1 indicate habitat preference; >1 habitat avoidance. bProbability that the mean ratio = 1.0.

Agriculture +++a +++ +++ +++ +++ +++ +++

62

Table 3.2. Habitat rankings based on pair-wise comparisons between habitat type distance ratios for second-order habitat selection, using seasonal home ranges of female bobcats monitored on Ichauway, Baker County, Georgia, 2001-2004. ____________________________________________________________________________________________________________

Agriculture Mixed pine- Pine Hardwood Wetland Shrub/Scrub Urban/Barren Pine hardwood regeneration ____________________________________________________________________________________________________________

Mixed ---a +b +++ +++ +++ +++ +++ pine-hardwood Pine --- --- + + + +++ +++ regeneration

Hardwood --- --- --- + + +++ +++

Wetland --- --- - - + +++ +++

Shrub/Scrub --- --- - - - + +

Urban/Barren --- --- --- --- --- - +

Pine --- --- --- --- --- - - ____________________________________________________________________________________________________________ a Three plus signs indicate row habitat significantly preferred over column and 3 minus signs indicate column habitat significantly preferred over row (t test, P>0.10). bA plus sign indicates that the row habitat type was closer to preference.

63

Table 3.3. Habitat type distance ratios for second-order selection using seasonal home ranges for male bobcats at Ichauway, Baker County, Georgia, 2001-2004. _________________________________________________________________________ Habitat type Meana t Pb

_________________________________________________________________________ Agriculture -0.2813 -3.00 0.0090

Shrub/Scrub 0.0257 0.21 0.8358

Hardwood -0.1601 -2.18 0.0460

Pine regeneration -0.1527 -1.13 0.2767

Pine -0.1374 -1.31 0.2100

Mixed pine-hardwood -0.0800 -1.32 0.2071

Wetland 0.0320 0.23 0.8250

Urban/Barren -0.0152 -0.19 0.8521 _________________________________________________________________________


Agriculture + + +++a +++ +++ +++ +++

64

Table 3.4. Habitat rankings based on pair-wise comparisons between habitat type distance ratios for second-order habitat selection, using seasonal home ranges of male bobcats monitored on Ichauway, Baker County, Georgia, 2001-2004. ____________________________________________________________________________________________________________

Agriculture Mixed pine- Pine Hardwood Urban/Barren Wetland Pine Shrub/Scrub hardwood regeneration ____________________________________________________________________________________________________________

Mixed - +b + +++ +++ +++ +++ pine-hardwood Pine ---a - + + + + +++ Hardwood --- - - + + +++ +++ Urban/Barren --- --- - - + + +++ Wetland --- --- - - - + +++ Pine --- --- - --- - - + regeneration Shrub/Scrub --- --- --- --- --- --- - ____________________________________________________________________________________________________________ a Three plus signs indicate row habitat significantly preferred over column and 3 minus signs indicate column habitat significantly preferred over row (t test, P>0.10). bA plus sign indicates that the row habitat type was closer to preference.

65

Table 3.5. Habitat type distance ratios for third-order selection using seasonal home ranges for male and female bobcats combined at Ichauway, Baker County, Georgia, 2001-2004. _________________________________________________________________________ Habitat type Meana t Pb

_________________________________________________________________________ Agriculture 0.0366 0.75 0.4561

Shrub/Scrub 0.0443 1.41 0.1673

Hardwood 0.0273 0.57 0.5748

Pine regeneration 0.1118 2.98 0.0048

Pine 0.1327 1.77 0.0844

Mixed pine-hardwood 0.2538 2.44 0.0190

Wetland -0.0004 -0.02 0.9875

Urban/Barren -0.0431 -2.01 0.0511 _________________________________________________________________________


Urban/Barren +b +++a + +++ +++ +++ +++

Shrub/Scrub --- - - - + + +++

Wetland - + + + +++ +++ +++

Hardwood ---a - + + +++ + +++

Agriculture - - - + + + +++

66

Table 3.6. Habitat rankings based on pair-wise comparisons between habitat type distance ratios for third-order habitat selection, using seasonal home ranges of all bobcats monitored on Ichauway, Baker County, Georgia, 2001-2004. ____________________________________________________________________________________________________________

Urban/Barren Wetland Hardwood Agriculture Shrub/Scrub Pine Pine Mixed pine- regeneration hardwood____________________________________________________________________________________________________________

Pine --- --- --- - - + + regeneration Pine --- --- - - - - + Mixed --- --- --- --- --- - - pine-hardwood ___________________________________________________________________________________________________________ a Three plus signs indicate row habitat significantly preferred over column and 3 minus signs indicate column habitat significantly preferred over row (t test, P>0.10). bA plus sign indicates that the row habitat type was closer to preference.

67

CHAPTER 4

BOBCAT DIETS IN A LONGLEAF PINE ECOSYSTEM

IN SOUTHWESTERN GEORGIA¹

_______________________ ¹Doughty, J., J. C. Cochrane, I. A. Godbois, L.M. Conner, and R. J. Warren. 2004. To be submitted to the Southeastern Naturalist.

68

ABSTRACT: Because northern bobwhite (quail; Colinus virginianus) hunting is an important

cultural and economic tradition in the Southeast and because quail management is an important

part of the longleaf pine (Pinus palustris)-wiregrass (Aristida stricta) ecosystem, it is important

to understand the impacts of bobcats (Lynx rufus) on quail. Here, we quantify seasonal and

annual diet of bobcats in a longleaf pine forest managed for quail. We collected 413 scats from

21 June 2001 to 20 June 2004. When possible, we identified prey items to species, but we

categorized scat components into 5 major prey groups (rodent, bird, rabbit, deer, and other) for

analysis. Bobcat diets varied among years (χ28 = 47.105, P < 0.001); therefore, we analyzed

seasonal variation for each year separately. Rodent comprised 75.1% of all scats, followed by:

other (21.8%), rabbit (Sylvilagus spp.; 20.6%), bird (13.1%) and deer (Odocoileus virginianus;

7.8%). Cotton rat (Sigmodon hispidus) comprised most (i.e., >60%) of the rodent category

during all years. Quail comprised 1.9% of all samples analyzed. Diet varied among seasons

during year 1 (χ212 = 19.934, P = 0.068) and year 2 (χ2

12 = 23.674, P = 0.023), but not during year

3 (χ212 = 17.352, P = 0.137). Bobcats were not a major predator of quail, but they were a major

predator of cotton rats and other rodents, which may compete with quail for resources. Further

investigation of the relationships between bobcats, rodents, and quail is necessary to better

understand whether bobcats may benefit quail by reducing populations of their competitors.

INTRODUCTION

Bobcats (Lynx rufus) primarily consume rabbits and hares, particularly cottontail rabbits

(Sylvilagus spp.), throughout their range (Anderson 1987). Rodents comprise the remaining bulk

of the diet, though species vary by habitat and locality. Some ground-dwelling avian (i.e., game

and nongame) and reptilian (i.e., snakes) species also are consumed.

69

In the Southeast, cotton rats (Sigmodon hispidus) and eastern cottontails (Sylvilagus floridanus)

compose most of the diet of bobcats during all seasons, and cotton rats replace eastern cottontails

as the primary prey source when they are more abundant (Beasom and Moore 1977, Miller and

Speake 1978, Boyle and Fendley 1987, Cochrane 2003).

Bobcats may exhibit a functional response to changing prey availability (Baker et al.

2001). White-tailed deer (Odocoileus virginianus) become an important food source in some

southern regions, especially during fall-winter when carrion from the hunting season is available,

and in late spring-summer, when fawns are available (Buttrey 1979, Story et al. 1982). Deer also

become an important food source during periods of low density of preferred prey (Beasom and

Moore 1977). In mountainous and highland regions of the South, squirrels (Sciurus spp.), pine

voles (Microtus pinetorum), and some bird species become important food sources (Buttrey

1979, Kitchings and Story 1979). Bobcats also ingest grass, either accidentally or intentionally

as a purgative (Miller and Speake 1978, Buttrey 1979).

Predators generally prey on species within certain size limits to optimize ease in capture

and energy returns (Rosenzweig 1966, McCord and Cordoza 1982). Bobcats concentrate kills on

prey from 150-5,500 g, and consume larger prey less frequently (Rosenzweig 1966, McCord and

Cordoza 1982, Story et al. 1982, Boyle and Fendley 1987, Anderson and Lovallo 2003). Sex

and age-related differences in food habits occur based on individual body size of bobcats. In

Arkansas, females consume more rats and mice than males (Fritts and Sealander 1978), and in

New Hampshire, males consume more white-tailed deer and fewer cottontails than females and

juveniles (Litvaitis et al. 1984). Such differences in food habits may decrease intraspecific

competition within bobcat populations (Rosenzweig 1966, Fritts and Sealander 1978).

70

Although bobcats historically have been considered a major predator of northern

bobwhite (quail; Colinus virginianus), few studies have specifically analyzed bobcat food habits

in areas managed for quail. On two quail plantations in southern Alabama, quail were not an

important part of bobcat diets despite a high density of quail (Miller and Speake 1978). On our

study area in southwestern Georgia, quail remains were found in 1.9% of all bobcat scats

collected over a 2-year period (Cochrane 2003). However, a different study found that quail

remains constituted between 0% and 12.5% of bobcat scats on one of two plantations

neighboring our study site (Schoch 2003).

Because quail hunting is an important cultural and economic tradition in the Southeast,

and quail management is an important part of the longleaf pine (Pinus palustris)-wiregrass

(Aristida stricta) ecosystem, it is important to understand the impacts of bobcats on quail (Burger

et al. 1999, Boring 2001). Our objective was to compare annual and seasonal diets of bobcats in

a longleaf pine forest managed for northern bobwhite. We predicted that diets would vary

among years, and that they would vary seasonally as a result of seasonal availability of prey

species.

STUDY AREA







71






















72















been harvested.

A variety of potential bobcat prey species are present at Ichauway. Small mammal

species include cotton rat, Eastern woodrat (Neotoma floridana), cotton mouse (Peromyscus

gossypinus), house mouse (Mus musculus), old-field mouse (P. polionotus), Eastern harvest

mouse (Reithrodontomys humulis), Southern short-tailed shrew (Blarina carolinensis) and least

shrew (Cryptotis parva) (Cochrane 2003). Eastern cottontail and marsh rabbit (S. palustris) are

present, and four species of sciurids are present (Eastern chipmunk [Tamias striatus], Southern

flying squirrel [Glaucomys volans], Eastern gray squirrel [Sciurus carolinensis], and fox squirrel

[S. niger].

73

METHODS

We collected scats on 30, 1-km long road transects once per month from June 2001-June

2004. Scat lines were assigned on secondary (frequently traveled, dirt roads) and tertiary (less

traveled, covered in grass) roads distributed throughout the study area. We also collected scats

opportunistically on the entire study area. Each sample was placed in a brown paper bag, labeled

with the date and location of collection, and frozen until processing. We collected ≥30 scats per

calendar season when possible (summer, fall, winter, spring).

Before processing, we thawed samples for 24 hours and oven-dried them for 72 hours at

60ºC (Baker et al. 1993, Griffin 2001). After weighing the samples, scats were sorted

macroscopically to separate and identify components (Baker et al. 1993). When possible, prey

items were identified to species using hair (Stains 1958), bone, teeth, and feathers (Baker et al.

1993). We categorized scat components into five major prey groups—rodent, bird, rabbit, deer,

and other (e.g., vegetation, opossum, raccoon, snake).

We calculated percent occurrence (i.e., frequency of occurrence divided by the total

number of scats examined within each season) for each prey category. We used a chi-squared

test of independence (Dowdy and Wearden 1991) in SAS (SAS Institute, Inc. 2003) to determine

if diet was dependent on year, and if diet varied seasonally within each year. We considered

statistical significance at α=0.10.

RESULTS

We collected 413 scats between 21 June 2001 and 20 June 2004 (n = 135 year 1, n = 130

year 2, n = 148 year 3). Bobcat diet varied among years (χ28 = 47.105, P < 0.001). The amount

of rodent and bird decreased from year 1 to year 2, and also decreased from year 2 to year 3.

74

Consumption of rabbit and other prey increased from year 1 to year 2, and from year 2 to year 3

(Figure 4.1). Deer decreased from year 1 to year 2, but increased from year 2 to year 3. Rodent

comprised 75.1% of all scats sorted, followed by: other (21.8%), rabbit (20.6%), bird (13.1%)

and deer (7.8%). Cotton rats comprised most (i.e., >60%) of the rodent category during all years

(year 1 = 74.0%; year 2 = 89.1%; year 3 = 61.6%). Northern bobwhite comprised 1.9% of all

samples analyzed (year 1 = 1.5%; year 2 = 2.3%; year 3 = 2.0%). Diet varied among seasons

during years 1 (χ212 = 19.934, P = 0.068; Figure 4.2) and 2 (χ2

12 = 23.674, P = 0.023; Figure 4.3),

but not during year 3 (χ212 = 17.352, P = 0.137; Figure 4.4). Species that comprised the other

category were: snake, raccoon, opossum, armadillo (Dasypus novemcinctus), skunk, bobcat, and

vegetation. For all years, vegetation (e.g., grass, seeds) was the most common food item in the

other category (year 1 = 37.5%; year 2 = 32.3%; year 3 = 86.1%).

DISCUSSION

Similar to other studies of bobcat diets in the southeastern U.S., bobcats in our study

primarily consumed rodents, rabbits, and other species (Davis 1955, Progulske 1955, Kight

1962, Beasom and Moore 1977, Miller and Speake 1978, Fritts and Sealander 1978, Kitchings

and Story 1979, Buttrey 1979, Fox and Fox 1982, Maehr and Brady 1986, Chamberlain and

Leopold 1999, Baker et al. 2001, Griffin 2001, Schoch 2003; Figure 4.5 and Table 4.1). Prey

selection is influenced by size, abundance, and energy returns associated with particular prey

species (Rosenzweig 1966, McCord and Cordoza 1982, Anderson and Lovallo 2003). Bobcats

are an opportunistic predator, but they may exhibit a functional response according to prey

availability and thus, are not purely opportunistic (McCord and Cordoza 1982, Baker et al. 2001,

Anderson and Lovallo 2003).

75

Among all years, rodent consumption decreased while rabbit consumption increased. An

overall increase in the rabbit population was a possible explanation for this occurrence;

however, we did not determine rodent or rabbit abundance during the study. Consumption of

prey in the other category also increased among all years, which may also reflect the

opportunistic nature of bobcat feeding habits. Vegetation was the most common food item in the

other category of bobcat prey species, probably ingested accidentally or intentionally as a

purgative (Miller and Speake 1978, Buttrey 1979).

During year 1, seasonal variation in bobcat diet was probably due to the increase in prey

consumption in the rabbit and other categories, particularly the increase from the winter to spring

seasons. Also, there were no rabbits consumed during the fall season, though rodent

consumption was highest during fall. Bird and deer were consumed more during the summer

season than any other seasons, possibly due to the presence of immature birds (nestlings and

fledglings), migratory bird species that are not present during other seasons, and fawns.

Seasonal variation during year 2 was likely due to the change in winter diet. While

rodent consumption declined and bird consumption was absent, deer and rabbit consumption

were higher than during any other season. Rabbit abundance may have increased during that

time period, and on Ichauway, rodent populations were generally lower during fall and winter

months (L.M. Conner, Joseph W. Jones Ecological Research Center, unpublished data).

There was no seasonal variation during year 3. Rodent consumption was lower overall

compared to other years, but it remained fairly constant during all seasons. Because overall bird

and deer consumption was low during summer, fall and winter, seasonal variation did not occur

despite a lack of consumption during the spring for both species. Rabbits and prey in the other

category fluctuated seasonally, but changes in consumption were not variable.

76

During all seasons and all years, rodents were the most common prey of bobcats on

Ichauway, which perhaps reflected the high rodent availability relative to other prey species

(Cochrane 2003). Management practices used for longleaf pine forest and quail management

may improve habitat for rodents (Cochrane 2003). On our study site, prescribed fire was a

primary land management tool, which increases and maintains a dense herbaceous understory

and early successional habitat, thereby providing abundant resources and habitat for prey

populations (Golley et al. 1965, Miller and Speake 1979). Planting agricultural crops and

maintaining quail food plots increases edge, which also provides ample resources for rodents

(Hall and Newsom 1976, Miller and Speake 1978).

Supplemental feeding increases prey abundance and concentrates prey (Boutin 1990).

Approximately 270 metric tons of grain sorghum are spread annually over 7,020 ha throughout

areas on our study site that are managed for quail between November and May (Godbois et al.,

2004). In a preliminary analysis of small mammal data collected at Ichauway, cotton rats were

5.5 more times abundant, cotton mouse were 1.5 times greater, Eastern harvest mouse were 2

times greater, and house mouse were 3.5 more times abundant in supplementally-fed versus

unfed areas (L.M. Conner, Joseph W. Jones Ecological Research Center, unpublished data).

Thus, current land management practices on Ichauway may have indirectly influenced bobcat

diets by providing abundant resources for rodent prey species, which resulted in more dense

rodent populations.

Contrary to other studies of bobcat diets in the Southeast, deer were consumed least

frequently during all years. Low deer consumption was probably influenced by the low density

of deer on the study area (4/km2), and because deer are more difficult to capture than rodents and

other prey species (Cochrane 2003).

77

In addition, deer may have been under-represented in scats, depending on length of exposure in

the field after deposition (Godbois et al. 2004b). Throughout the study, birds also were not a

major prey item. Fifty-four out of 413 scat samples contained bird remains, and only eight of the

bird remains were quail. Our findings were similar to Miller and Speake (1978), in which quail

comprised only two of 218 bobcat scats sampled from two quail plantations in Alabama. In

contrast, Schoch (2003) found quail remains (i.e., feathers or eggs) in seven of 66 bobcat

stomachs collected on a quail plantation in Georgia. Similar to our results, rodents were the

primary prey items for both of these studies, despite high densities of quail on the study areas.

Bobcats were not a primary predator of quail in our study. Although land management

practices such as prescribed fire, crop planting, maintenance of food plots, and supplemental

feeding are beneficial to quail, they also attract rodents and other species (Stoddard 1931).

Snakes and birds of prey, other potential quail predators, may be attracted to areas with dense

rodent populations. Cotton rats may compete with quail for food sources directly, and they

damage plants used by quail (Simpson 1976). Cotton rats also destroy northern bobwhite nests

(Stoddard 1931, Simpson 1976, Staller 2001). At Ichauway, bobcats were 10 times closer to

supplemental food than expected (Godbois et al. 2004a). Thus, in areas where densities of

cotton rats and other rodents are high, such as supplemental-feed areas and field edges, bobcats

may benefit quail by reducing populations of their competitors.

Further investigation of the relationships between bobcats, rodents, and quail is

necessary. Bobcat diet and prey abundance should be studied concurrently, to better explain

fluctuations in prey consumption annually and seasonally. Future studies of bobcat diet at

Ichauway should incorporate analysis using stomach or intestinal contents to better quantify the

presence of prey items that may be underrepresented in scat alone (e.g., deer and egg fragments).

78

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Kitchings, J. T., and J. D. Story. 1979. Home range and diet of bobcats in eastern Tennessee.

Bobcat Research Conference Proceedings. National Wildlife Federation Scientific and

Technical Series 6:47-52.

81



Litvaitis, J. A., C. L. Stevens, and W. W. Mautz. 1984. Age, sex, and weight of bobcats in

relation to winter diets. Journal of Wildlife Management 48:632-635.

Maehr, D. S, and J. R. Brady. 1986. Food habits of bobcats in Florida. Journal of Mammalogy

67:133-138.

McCord, C. M. and J. E. Cordoza. 1982. Bobcat and lynx. Pages 728-766 in J. A. Chapman

and G. A. Feldhamer, eds. Wild Mammals of North America. Johns Hopkins University

Press, Baltimore, Maryland, USA.

Miller, S. D., and D. W. Speake. 1978. Prey utilization on quail plantations in southern

Alabama. Proceedings of Southeastern Association of Fish and Wildlife Agencies

32:100-111.




Series 6.

Progulske, D. R. 1955. Game animals utilized as food by bobcat in the southern Appalachians.

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Rosenzweig, M. L. 1966. Community structure of sympatric Carnivora. Journal of




82

Schoch, B. N. 2003. Diet, age, and reproduction of mesomammalian predators in response to

intensive removal during the quail nesting season. M. S. Thesis. University of Georgia,

Athens.

Simpson, R. C. 1976. Certain aspects of bobwhite quail’s life history and population dynamics

in southwest Georgia. Technical Bulletin. Georgia Department of Natural Resources,

Atlanta, Georgia. 117pp.

Stains, H. J. 1958. Field key to guard hair of middle western furbearers. Journal of Wildlife

Management. 22:95-97.

Staller, E. L. 2001. Identifying predators and fates of northern bobwhite nests using miniature

video camera. M. S. Thesis. University of Georgia, Athens.

Stoddard, H. L. 1931. The Bobwhite Quail: Its Habits, Preservation and Increase

Charles Scribner’s Sons, New York. 559pp.

Story, J. D., W. J. Galbraith, and J. T. Kitchings. 1982. Food habits of bobcats in eastern

Tennessee. Journal of Tennessee Academic Science 57:29-32.

83

Figure 4.1. Annual percent occurrence of prey items consumed by bobcats based on scat

analysis on Ichauway, Baker County, Georgia, 2001-2004.

84

0

10

20

30

40

50

60

70

80

90

100

rodent bird deer rabbit other

prey category

% sc

at c

onta

inin

g ca

tego

ry

year 1year 2year 3

85

Figure 4.2. Seasonal percent occurrence of prey items consumed by bobcats based on analysis of

135 scats on Ichauway, Baker County, Georgia, 2001-2002.

86

0

10

20

30

40

50

60

70

80

90

100


prey category

% sc

at c

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g ca

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summerfallwinterspring

87



88

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10

20

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40

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60

70

80

90

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prey category

% sc

at c

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89



90

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10

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prey category

% sc

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91

Figure 4.5. Seasonal percent occurrence of prey items consumed by bobcats (S01=Summer

2001; F01=Fall 2001; W02=Winter 2002; S02=Spring 2002; Su02=Summer 2002; F02=Fall

2002; W03=Winter 2003; S03=Spring 2003; Su03=Summer 2003; F03=Fall 2003; W04=Winter

2004; S04=Spring 2004) based on analysis of 413 scats on Ichauway, Baker County, Georgia,

2001-2004.

92

0

10

20

30

40

50

60

70

80

90

100

Su01 F01 W02 S02 Su02 F02 W03 S03 Su03 F03 W04 S04

season

% sc

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g ca

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rodentbirddeerrabbitother

93

Table 4.1. Studies documenting bobcat diet by percent occurrence in the southeastern United States, using the gastrointestinal tract (GI; stomach or intestines) or scat for analysis. ______________________________________________________________________________ Reference State n Rodent Bird Deer Rabbit Other Analysis Technique ______________________________________________________________________________

Davis,1955 AL 239 18.8 6.9 10.5 63.2 20.6 GIb

Progulske, 1955 VA 124 66.9 14.9 9.9 54.6 62.8 Scat

Kight, 1962 SC 317 65.6 21.8 0.0 43.8 29.0 Scat

Beasom and Moore, 1977 TX 125 96.5 18.5 3.0 23.0 13.0 GI

Fritts and Sealander, 1978 AR 150 21.0 8.0 7.0 39.0 52.0 GI

Miller and Speake, 1978 AL 273 49.1 10.0 9.5 29.3 38.5 GI

Miller and Speake, 1978 AL 218 87.2 15.6 1.4 37.6 43.6 Scat

Buttrey, 1979 TN 49 60.0 12.2 20.4 34.7 36.6 GI/Scat

Kitchings and Story, 1979 TN 31 46.5 7.0 19.5 68.5 46.5 Scat

Fox and Fox, 1982 WV 172 44.1 8.9 50.1 23.6 5.5 GI

Maehr and Brady, 1986 FL 413 42.4 18.7 2.4 85.5 25.7 GI

Chamberlain and Leopold, 1999a MS 591 21.6 15.3 28.2 13.0 Scat Baker et al., 2001 GA 357 15.7 7.8 36.4 46.2 13.2 Scat Griffin, 2001 SC 179 43.0 13.7 19.0 16.0 8.7 Scat Schoch, 2003 GA 66 62.1 10.6 9.1 28.8 18.2 GI This study GA 413 75.1 13.1 7.8 20.6 21.8 Scat ______________________________________________________________________________ aBird included in 'other' category.

94

CHAPTER 5

CONCLUSIONS AND MANAGEMENT IMPLICATIONS

95

Bobcats (Lynx rufus) are considered an apex predator in certain forested ecosystems of

the southeastern U.S., including the longleaf pine (Pinus palustris)-wiregrass (Aristida stricta)

ecosystem of southwestern Georgia (Conner et al. 2000). However, control of bobcats and other

mammalian predators is often suggested as a method to increase game bird species abundance.

Although bobcats have historically been considered a major predator of northern bobwhite

(quail; Colinus virginianus), few studies have specifically analyzed bobcat food habits in areas

managed for quail. Because quail hunting is an important cultural and economic tradition in the

Southeast, and quail management is an important part of the longleaf pine ecosystem, it is

important to understand the impacts of bobcats on quail (Boring 2001, Burger et al. 1999). Thus,

one of our main objectives was to quantify seasonal diets of bobcats in a longleaf pine forest

managed for northern bobwhite. Our other objectives were to determine if annual and seasonal

bobcat home ranges varied by season and sex, determine habitat use at 2 spatial scales and

compare habitat use between the sexes and among seasons, and determine whether activity status

and time-of-day influenced bobcat habitat use.

Only 8 out of 413 scats collected over 3 years contained quail remains. Although we

could not account for quail eggs due to the type of diet analysis technique we used, we concluded

that bobcats were not a major predator of quail. During all seasons and all years, rodents were

the most common prey of bobcats on Ichauway, likely due to high rodent abundance relative to

other prey species (Cochrane 2003). Management practices used for longleaf pine forest and

quail management may improve habitat for rodents. On our study site, prescribed fire was a

primary land management tool, which increases and maintains a dense herbaceous understory

and early successional habitat, thereby providing abundant resources and habitat for prey

populations (Golley et al. 1965, Miller and Speake 1979).

96

Planting agricultural crops and maintaining quail food plots increases edge, which also provides

ample resources for rodents (Hall and Newsom 1976, Miller and Speake 1978). Supplemental

feeding may concentrate rodent populations, and bobcats at Ichauway were 10 times closer than

expected to supplementally-fed versus non-fed areas (Godbois et al. 2004).

Although home ranges for male bobcats were larger than for females on our study site,

home ranges of both sexes were smaller than home ranges previously reported for the

southeastern United States (Buie et al. 1979, Kitchings and Story 1979, Hamilton 1982, Shiftlet

1984, Lancia et al. 1986, Rucker et al. 1989, Conner et al. 1992, Conner et al. 2001). Land

management practices such as prescribed burning and maintenance of food plots contributed to

high-quality habitat with ample prey, which likely contributed to smaller home range sizes of

bobcats on our study area. Female home range sizes varied seasonally. Between winter and

spring during all 3 years of the study, female home range sizes declined, which was likely due to

females denning and restricting their movements (Bailey 1974, Knick 1990). The smallest

female home range occurred during Summer 2002, which corresponded with the post-parturition

period for that year. Females probably restricted their movements to provide prey to their kittens

(Bailey 1979, Jackson and Jacobson 1987, Conner et al.1992). Home ranges increased between

summer and fall seasons, during which kittens become old enough to travel with their mother

(Bailey 1979). Seasonal variation in bobcat home range size also may be influenced by prey

availability and breeding behavior.

In the longleaf pine ecosystem of Ichauway, bobcats selected habitats to include in their

home range (Johnson's second order of selection), but did not select habitats within the home

range (Johnson's third order of selection). At Johnson's second order, female bobcats were closer

to all habitat types than expected.

97

Males were closer then expected to all habitat types except shrub/scrub and wetland, suggesting

a preference for edges (Conner et al. 2003). Thus, bobcats appeared to prefer edge when

establishing home ranges. Edge provides travel routes and access to hunting areas such as

agriculture fields, and it is typically where an abundance of prey is found (Landers and Mueller

1986).

Similar to other studies in the Southeast, our bobcats preferred agriculture (Cochrane

2003, Conner et al. 1992), mixed pine-hardwood (Cochrane 2003), pine (Conner et al. 1992), and

hardwood habitats (Hall and Newsom 1976, Lancia et al. 1986). Bobcats likely selected early

and mid-successional habitats due to prey abundance and availability. Cotton rats (Sigmodon

hispidus) and eastern cottontails (Sylvilagus floridanus), 2 primary bobcat prey species in the

Southeast, are most abundant in dense areas of early to mid-successional grass/forb-shrub

vegetation (Boyle and Fendley 1987). Agricultural areas also attract rodent and lagomorph

species (Cummings and Vessey 1994). Hardwood and mixed pine-hardwood habitats contain a

dense herbaceous understory and shrub interspersion necessary to provide essential cover for

prey species, and such habitats also provide bobcats with cover (Golley et al. 1965, Schnell

1968).

We expected active animals to select agriculture and other early-to-mid-successional

habitat types more than inactive animals. Because bobcats are considered a nocturnal and/or

crepuscular species, we also expected bobcats to prefer early-to-mid-successional habitats during

the nighttime and crepuscular periods. Activity at crepuscular time periods coincides with

activity peaks for rodents and lagomorphs, which are primary bobcat prey.

98

Contrary to our hypotheses, bobcats did not select habitat differently according to activity status

or time-of-day. Because bobcats selected early-to-mid-successional habitat types to include in

their home ranges and such habitats may provide dense herbaceous cover, resting animals (i.e.,

inactive) likely did not have to find cover in other habitat types. If preferred habitats provided

ample prey and resting sites, then bobcats probably did not have to move to other habitats during

periods of rest or at different times throughout the diel period.

We suggest that bobcat habitat selection is probably determined by prey availability.

Habitat should be managed to provide a dense herbaceous layer for prey and bobcats. Prescribed

fire can accomplish this goal within the longleaf pine ecosystem. In addition, managing for a

mosaic of diverse habitat types increases the amount of edge available, which is an important

component of bobcat habitat.

Bobcat den sites have been discovered in hollow logs, rocky outcrops (Gashwiler et al.

1961, Cochrane 2003), at the base of tree stumps in timber-harvested areas (Kitchings and Story

1984), and in thickets and brush piles (Anderson 1987, Cochrane 2003). Den sites in human-

made structures, such as abandoned buildings, also have been observed (Bailey 1974). Three of

the 5 dens located were in bulldozed piles of trees and brush, by-products of human activities.

Den site selection is probably determined by protection from environmental conditions, and

possibly prey availability, since females are limited to hunting prey in close proximity to

unprotected kittens (Bailey 1979). Prescribed burning may create thickets and hollow stumps,

thereby increasing the availability of den sites for bobcats (Young 1958, Kitchings and Story

1984).

99

During the study, 14 confirmed bobcat mortalities occurred. Eight of the mortalities were

human-related (5 trapped and/or killed off-site; 3 from vehicle collisions). One bobcat was

killed by another felid, and 4 bobcats died from unknown causes.

Similar to studies of bobcats in other ecosystems of the southeastern U.S., bobcat home

range size and habitat use within the longleaf pine ecosystem were influenced by prey

availability and abundance. Although bobcats were not a major predator of quail, further

investigation of the relationships between bobcats, rodents, and quail is necessary. Bobcat diet

and prey abundance should be studied concurrently, to better explain fluctuations in prey

consumption annually and seasonally. Future studies of bobcat diet at Ichauway should

incorporate analysis using scat and stomach or intestinal contents to better quantify the presence

of particular prey items that may be underrepresented in scat alone (e.g., deer and egg

fragments).

Although our results suggest that prescribed burning and other quail management

practices may impact bobcat ecology, future research on Ichauway should document prey

population densities and bobcat locations in burned versus non-burned areas and in food plots

versus non-food plots, which might provide stronger evidence of the role land management

practices play in bobcat ecology in the longleaf pine forest.

Literature Cited





100

_____. 1979. Den ecology, population parameters, and diet of eastern Idaho bobcats. Pages 62-


conference. National Wildlife Federation of Scientific and Technical Series 6.






Reports 82(10.147).








Thesis, University of Georgia, Athens, Georgia, USA. 87 pp.




101

_____, B. D. Leopold, and M. J. Chamberlain. 2000. Multivariate habitat models for bobcats in

southern forested landscapes. Pages 51-55 in Proceedings of a symposium on current

bobcat research and implications for management. The Wildlife Society 2000


_____ and B. D. Leopold. 2001. Spatio-Temporal relationships among adult bobcats in central

Mississippi. Pages 45-50 In: Woolf, A., C. K. Nielsen, and R. D. Bluett, (eds.).

Proceedings of the Symposium on Current Bobcat Research and Implications for

Management. The Wildlife Society 2000 Conference, Nashville, Tennessee, USA.

_____, M. D. Smith, and L. W. Burger. 2003. A comparison of distance-based and

classification-based analyses of habitat use. Ecology 84:526-531.

Cummings, J. R. and S. H. Vessey. 1994. Agricultural influences of movement patterns of

white-footed mice (Peromyscus leucopus). American Midland Naturalist 132:209-218.

Gashwiler, J. S., W. L. Robinette, and O. W. Morris. 1961. Breeding habits of bobcats in Utah.




518.




bottomland hardwoods of southern Louisiana. Proceedings of the Annual Conference

Southeastern Association of Fish and Wildlife Agencies 30: 427-436.

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Hamilton, D. A. 1982. Ecology of the bobcat in Missouri. Thesis, University of Missouri,

Columbia, Missouri, USA. 152 pp.


managed southern forest ecosystems. Final Report Federal Aid Project, W-48-30, 31, 32,

33, 32. Mississippi Department of Wildlife Conservation Study XX. 69 pp.

Kitchings, J. T. and J. D. Story. 1979. Home range and diets of adult bobcats in eastern

Tennessee. Pages 47- 52 in L. G. Blum and P. C. Escherich, eds. Proceedings of the

bobcat research conference. National Wildlife Federation Scientific and Technical Series

6.




Idaho. Wildlife Monographs 108. 42 pp.


habitat use by bobcats on Croatan National Forest, North Carolina. Pages 425-436 in:

S. D. Miller and D. D. Everett, (eds.). Cats of the world: biology, conservation, and

management. Caesar Kleberg Wildlife Research Institute, Kingsville, Texas, USA.

Landers, J. L. and B. S. Mueller. 1986. Bobwhite quail management: a habitat approach.

Tall Timbers Research Station and Quail Unlimited, Tallahassee, Florida, USA.


Alabama. Proceedings of the Annual Conference of the Southeastern Association of Fish

and Wildlife Agencies 32:100-111.

103




Series 6.





Shiftlet, B. L. 1984. Movements, activity, and habitat use of the bobcat in upland mixed pine-

hardwoods. Thesis, Louisiana State University, Baton Rouge, Louisiana, USA.


193 pp.

104

APPENDIX A

MANAGEMENT ZONES ON ICHAUWAY, BAKER COUNTY, GEORGIA, 2000-2004

105

Conservation zones (40% of property) are managed with prescribed burning for longleaf pine

restoration. Multiple-use zones (60% of property) are managed with prescribed burning,

maintenance of food plots, supplemental feeding, and some predator control for northern

bobwhite.

106

APPENDIX B

MORPHOLOGICAL DATA COLLECTED FOR 50 ADULT BOBCATS

CAPTURED AND RADIO-COLLARED BETWEEN

DECEMBER 2000 AND MAY 2004, ICHAUWAY, BAKER COUNTY, GEORGIA

107

Morphological data collected for 50 adult bobcats captured and radio-collared between December 2000 and May 2004, Ichauway, Baker County, Georgia. ______________________________________________________________________________ Date Cat # Sexa Wtb Total Tail Hind Front (kg) length (cm) length (cm) foot length of ear (cm) length (cm) ______________________________________________________________________________

12/13/00 01 F 5.75 860 150 165 65 01/10/01 04 F 6.75 902 143 160 61 01/24/01 10 F 7.33 902 152 164 59 01/25/01 12 M 7.00 932 125 160 58 02/07/01 15 F 6.25 917 138 154 54 02/07/01 16 M 9.50 941 132 170 62 03/08/01 18 F 6.75 890 135 165 60 03/27/01 23 F 7.25 912 121 168 66 04/08/01 25 F 7.75 880 130 170 70 04/20/01 27 F 7.00 915 131 170 64 04/24/01 28 F 6.75 890 138 161 62 05/01/01 30 M 7.00 954 142 179 62 06/03/01 34 M 8.10 1000 170 170 70 06/04/01 05 M 5.60 940 160 160 60 11/16/01 36 M 10.50 1040 160 185 70 12/07/01 37 F 6.50 863 113 150 62 12/19/01 39 F 7.83 850 130 165 70 12/26/01 41 M 10.00 1010 140 175 70 03/05/02 06 F 7.40 940 145 160 60 03/05/02 45 M 8.50 1040 150 170 70 03/06/02 47 F 7.00 850 120 160 60 03/07/02 40 F 6.50 905 145 160 65 03/09/02 50 M 7.25 985 160 170 65 03/26/02 46 M 8.45 970 145 165 60 04/30/02 48 F 6.50 870 140 153 68 05/15/02 42 F 6.25 895 140 160 60 06/21/02 51 F 7.25 940 155 170 65 12/04/02 49 F 6.30 860 121 164 57 12/12/02 54 F 6.20 900 130 165 51 12/13/02 57 M 10.20 985 165 180 49 01/30/03 60 F 8.75 844 137 173 52 01/30/03 61 M 6.75 985 145 182 55 02/17/03 65 M 9.95 940 140 173 58 03/04/03 67 F 5.50 908 135 165 51 _____________________________________________________________________________

108

______________________________________________________________________________

Date Cat # Sexa Wtb Total Tail Hind Front (kg) length (cm) length (cm) foot length of ear (cm) length (cm) ______________________________________________________________________________

04/25/03 71 M 9.95 995 155 175 55 05/21/03 72 F 5.50 865 140 155 55 12/16/03 75 M 6.75 964 152 172 57 12/20/03 76 F 6.50 898 138 162 51 01/11/04 31 M 11.3 1047 180 180 64 01/21/04 80 F 6.55 935 152 175 62 01/28/04 82 F 6.50 870 152 164 70 02/01/04 83 M 10.30 985 140 179 70 03/11/04 11 M 7.50 966 166 174 58 03/18/04 53 M 8.75 1019 177 173 70 04/13/04 44 F 6.00 884 119 169 74 04/21/04 89 F 6.75 900 110 158 62 05/08/04 86 F 5.75 910 142 161 65 05/14/04 66 F 5.00 860 138 155 68 05/27/04 77 F 5.40 866 140 161 55 05/30/04 90 F 7.75 912 116 154 58 ______________________________________________________________________________ aF = Female; M = Male. bWeight.

109

APPENDIX C

ANNUAL AND SEASONAL

ADAPTIVE KERNEL (ADK) AND MINIMUM CONVEX POLYGON (MCP)

HOME RANGE SIZE ESTIMATES FOR BOBCATS ON

ICHAUWAY, BAKER COUNTY, GEORGIA, 2001-2004

110

Mean seasonal and annual home range sizes (km2) for 44 bobcats on Ichauway, Baker County, Georgia, 2001-2004, according to sex and home range estimator. ______________________________________________________________________________ Season/Year ADKa Estimate MCPb Estimate ______________ ______________ M F M F ______________________________________________________________________________

Fall 2001 2.1 5.1 2.6 3.0

Winter 2002 15.9 4.0 4.0 2.5

Spring 2002 8.1 3.3 4.4 1.8

Summer 2002 3.7 2.8 6.7 1.8

Fall 2002 7.2 5.6 4.9 3.3

Winter 2003 11.1 8.5 6.6 4.5

Spring 2003 10.5 4.5 6.6 2.8

Summer 2003 7.9 6.1 4.8 3.9

Fall 2003 6.6 5.9 4.0 3.6

Winter 2004 11.3 7.1 6.9 4.2

Spring 2004 9.6 5.6 6.5 3.7

All Seasonsc 8.5 5.3 5.3 3.2

Annual 2002-2003d 10.0 4.3 7.1 3.3

Annual 2003-2004d 12.3 8.1 9.7 6.9 All Yearse 11.2 6.2 8.4 5.1 ______________________________________________________________________________

a95% Adaptive Kernel. b95% Minimum Convex Polygon. cMean home range for all seasons pooled. dMean annual home range from Spring 2002-Winter 2003 and Spring 2003-Winter 2004. eMean annual home range for both years pooled.

111

Individual 95% adaptive kernel and 95% minimum convex polygon annual home range estimates (km2) for 44 bobcats on Ichauway, Baker County, Georgia, 2001-2004 (a minimum of 30 locations/bobcat/season for 4 consecutive seasons were obtained for analysis) ______________________________________________________________________________ Cat # Sexa ADKb Estimate MCPc Estimate __________________ __________________ Year 1d Year 2e Year 1 Year 2 ______________________________________________________________________________

01 F 2.9 2.2 04 F 4.0 3.3 10 F 5.2 4.3 23 F 4.4 3.4 36 M 9.8 7.7 41 M 13.3 10.1 15 F 3.1 6.8 2.3 5.2 18 F 6.1 7.2 44.8 5.9 25 F 6.5 9.7 4.2 7.1 34 M 12.0 16.1 7.3 12.8 37 F 2.9 5.3 2.7 3.6 39 F 4.7 7.9 3.4 5.8 45 M 5.0 7.6 3.5 5.3 47 F 3.4 4.7 2.6 3.6 06 F 5.0 3.8 42 F 6.5 5.3 49 F 5.5 4.7 60 F 25.0 26.7 63 F 4.9 3.7 67 F 8.2 6.6 71 M 13.4 10.9 ______________________________________________________________________________ aF = Female; M = Male b95% Adaptive Kernel. cMinimum Convex Polygon. dMean annual home range from Spring 2002-Winter 2003. eMean annual home range from Spring 2003-Winter 2004.

112

Individual 95% adaptive kernel seasonal home range estimates (km2; F01=Fall 2001; W02=Winter 2002; S02=Spring 2002; Su02=Summer 2002; F02=Fall 2002; W03=Winter 2003; S03=Spring 2003; Su03=Summer 2003; F03=Fall 2003; W04=Winter 2004; S04=Spring 2004) for 44 bobcats on Ichauway, Baker County, Georgia, 2001-2004 (a minimum of 30 locations/bobcat/season were obtained for analysis) ______________________________________________________________________________ Cat # Sex F01 W02 S02 Su02 F02 W03 S03 Su03 F03 W04 S04 ______________________________________________________________________________ 01 F 5.0 1.4 1.2 1.8 3.9 5.9 4.3 5.4 04 F 3.5 3.1 4.0 3.5 4.3 4.2 7.6 4.3 05 M 4.6 3.1 3.0 06 F 3.1 3.6 17.5 3.1 2.9 4.2 5.1 5.5 10 F 3.9 4.3 1.2 3.2 2.7 6.1 7.7 13.8 7.8 11 M 10.7 15 F 2.8 1.6 2.7 1.2 3.8 3.4 3.0 4.5 5.2 9.6 7.2 16 M 5.6 2.8 3.4 4.3 5.1 18 F 4.6 1.6 3.8 3.9 5.6 14.0 4.9 6.7 4.7 7.1 3.4 23 F 4.5 4.8 5.5 2.0 3.7 9.5 3.0 9.9 7.9 25 F 1.2 1.7 3.7 1.3 6.6 6.3 7.2 12.3 8.3 8.3 6.1 26 F 2.2 27 F 18.4 14.2 1.0 6.7 7.4 30 M 3.2 80.1 9.8 8.8 8.5 9.0 31 M 5.9 6.4 34 M 5.4 5.7 6.8 4.1 4.3 17.5 10.7 9.8 9.0 17.6 19.4 36 M 1.2 5.2 6.8 9.8 8.3 7.9 5.9 8.7 37 F 2.1 2.7 2.0 1.7 2.7 1.5 3.5 4.4 5.4 6.0 39 F 5.7 5.4 2.6 2.4 6.7 4.2 6.7 7.7 6.8 4.0 40 F 6.0 31.0 34.7 7.8 41 M 6.2 3.1 6.2 11.9 13.0 13.5 9.0 6.0 42 F 3.0 5.7 5.3 3.2 4.3 6.5 6.6 6.8 44 F 3.5 45 M 2.4 3.7 3.1 4.3 8.8 8.5 5.3 5.8 5.2 46 M 0.7 47 F 3.3 2.0 1.7 4.2 3.1 3.3 5.9 5.1 3.2 48 F 6.1 49 F 3.1 3.1 7.4 4.2 5.3 3.4 51 F 0.8 3.6 53 M 7.2 57 M 16.4 9.1 7.4 ______________________________________________________________________________

113

______________________________________________________________________________ Cat # Sexa F01 W02 S02 Su02 F02 W03 S03 Su03 F03 W04 S04 ______________________________________________________________________________ 60 F 3.9 6.9 8.1 5.9 9.3 10.2 61 M 10.1 9.2 6.9 63 F 6.7 3.6 3.7 2.4 3.1 65 M 6.5 3.5 66 F 4.9 67 F 3.1 4.5 6.0 7.7 5.5 71 M 13.4 8.5 8.7 15.8 10.4 75 M 19.5 80 F 3.0 2.0 82 F 8.9 9.7 83 M 4.2 3.7 86 F 5.7 89 F 10.3 ______________________________________________________________________________ aF = Female; M = Male

114

Individual 95% minimum convex polygon seasonal home range estimates (km2; F01=Fall 2001; W02=Winter 2002; S02=Spring 2002; Su02=Summer 2002; F02=Fall 2002; W03=Winter 2003; S03=Spring 2003; Su03=Summer 2003; F03=Fall 2003; W04=Winter 2004; S04=Spring 2004) for 44 bobcats on Ichauway, Baker County, Georgia, 2001-2004 (a minimum of 30 locations/bobcat/season were obtained for analysis) ______________________________________________________________________________ Cat # Sex F01 W02 S02 Su02 F02 W03 S03 Su03 F03 W04 S04 ______________________________________________________________________________ 1 F 2.5 0.8 0.7 1.3 2.0 2.8 2.5 2.0 4 F 2.0 1.6 2.3 2.4 2.5 2.5 2.7 4.2 3.5 5 M 2.6 1.3 2.1 6 F 1.5 2.0 7.7 1.9 1.8 2.6 3.6 3.9 10 F 2.5 2.6 0.6 1.9 1.7 3.5 6.0 13.8 7.8 11 M 8.3 15 F 1.8 1.1 1.4 0.8 2.2 2.0 1.9 3.2 3.2 5.5 4.4 16 M 3.6 1.8 1.6 24.3 3.2 18 F 2.8 1.0 2.3 2.5 3.0 5.7 3.0 4.4 2.4 3.7 2.5 23 F 3.1 3.3 3.8 1.3 2.4 3.0 5.7 7.0 4.6 25 F 0.8 1.0 1.7 1.1 3.7 4.0 4.1 6.8 5.3 4.6 2.8 26 F 1.3 27 F 10.7 9.3 0.4 4.4 4.7 30 M 2.2 14.7 5.6 38.3 5.6 5.6 5.2 31 M 2.8 4.4 34 M 3.8 3.9 5.4 3.2 3.6 9.9 7.4 6.1 6.5 14.5 13.9 36 M 0.7 3.2 6.3 6.9 5.1 4.8 3.7 5.5 37 F 1.1 1.8 1.4 1.4 1.8 1.1 2.6 2.8 3.3 3.2 39 F 3.5 2.9 1.7 1.4 4.4 2.7 4.1 4.4 3.9 2.2 40 F 4.1 15.8 18.2 5.6 41 M 3.8 2.5 4.8 7.7 8.5 8.2 5.7 3.7 42 F 1.6 3.3 2.5 1.9 2.4 3.6 5.3 3.7 44 F 1.9 45 M 1.3 1.7 2.2 2.5 7.5 4.3 3.2 3.1 3.0 46 M 0.4 47 F 1.4 1.4 1.4 2.5 2.0 2.0 3.7 3.1 2.2 48 F 1.9 49 F 2.7 1.7 4.6 3.1 3.2 2.5 51 F 0.5 2.9 53 M 4.8 57 M 9.4 4.6 3.9 ______________________________________________________________________________

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______________________________________________________________________________ Cat # Sex F01 W02 S02 Su02 F02 W03 S03 Su03 F03 W04 S04 ______________________________________________________________________________ 60 F 2.3 4.1 4.9 4.0 6.2 7.7 61 M 5.4 4.9 3.9 63 F 3.3 2.5 2.7 1.7 2.1 65 M 4.4 1.5 66 F 3.8 67 F 1.8 2.9 4.0 5.1 3.6 71 M 8.0 6.2 6.0 8.4 8.6 75 M 8.5 80 F 1.4 1.2 82 F 4.5 5.4 83 M 4.6 1.6 86 F 2.8 89 F 7.4 ______________________________________________________________________________ aF = Female; M = Male

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APPENDIX D

DESCRIPTION OF FIVE BOBCAT DENS LOCATED IN A LONGLEAF PINE

ECOSYSTEM, ICHAUWAY, BAKER COUNTY, GEORGIA, 2002-2004

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We confirmed location of five dens through the duration of the study. The first den was

located 17 April 2002 in a bulldozed pile of trees, stumps, and branches. The den was audibly

confirmed by the sound of least 2 kittens. The mother was observed in the den. Den 2 was

located 17 April 2003, in a brush pile bulldozed for logging, surrounded by briars and sassafras

(Sassafras spp.). At least two kittens were heard, but there was no visual confirmation of kittens

or adult. The third den was located 24 April 2003 in a hollowed-out water oak tree (Quercus

nigra) in a hardwood bottom close to the Flint River. Two kittens with closed eyes were

observed with the adult female. Den 4 was located 24 April 2003 in a fallen-down hollow log

surrounded by dense vegetation. At least 2 kittens were heard, and the adult female was

observed fleeing the den. The fifth den was located 30 April 2004 in a brush pile created as a

by-product of logging near a primary road. At least 1 kitten was heard, but there was no visual

confirmation of the kittens or mother.

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APPENDIX E

DESCRIPTION OF BOBCAT MORTALITIES IN SOUTHWESTERN GEORGIA, 2001-2004

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Mortality events for bobcats captured on Ichauway, Baker County, Georgia, 2001-2004 ____________________________________________________________________________________________________________ Cat # Sexa Ageb Date of initial Date of last Date of mortality Cause of Mortality capture radio-location ____________________________________________________________________________________________________________

07 F A 01/22/01 06/19/01 06/21/01 vehicle collision 14 M A 02/02/01 10/05/01 10/29/01 trapped off-site (LL Plantationc) 19 M A 03/14/01 N/Ad 05/24/02 unknown; dead in trap 03 M A 01/04/01 never relocated 09/27/01 trapped off-site (PB Plantatione) 26 M A 04/19/01 01/27/02 01/28/02 killed by another felid 32 M A 05/16/01 05/19/01 06/28/02 killed off-site (private plantation) 51 F A 06/21/02 10/27/02 12/3/02 unknown 54 F A 12/12/02 never relocated 12/30/02 unknown 16 M A 02/07/01 02/18/03 03/04/03 unknown - possibly vehicle collision 30 M A 05/21/01 07/02/03 07/08/03 unknown 36 M A 11/16/01 11/26/03 11/29/03 killed off-site (LL Plantation) 41 M A 01/26/02 12/19/03 12/26-12/28/03 vehicle collision 73 M J 05/30/03 N/A 01/28/04 trapped off-site (LL Plantation) 75 M A 12/16/03 04/14/04 04/29/04 probably trap injury; broken neck 83 M A 02/01/04 05/28/04 05/29/04 vehicle collision ____________________________________________________________________________________________________________ aF = Female; M = Male. bA = Adult; J = Juvenile. cLongleaf Plantation, located 3-4 miles away from Ichauway. dNot applicable to juvenile bobcats (not radio-collared) or bobcats recaptured as adults and not radio-collared during first capture.

ePinebloom Plantation, located >10 miles away from Ichauway.

bobcat (lynx rufus · bobcat (lynx rufus) ecology in a longleaf pine ecosystem in southwestern...

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