• TABLE OF CONTENTS
HIDE
 Title Page
 Acknowledgement
 Table of Contents
 Executive summary
 Preface
 Spatial distributions of bobcats...
 Scent-station indices as indicators...
 Appendix A: Capture date, sex,...
 Appendix B: Capture date, sex,...






Group Title: Florida Cooperative Fish and Wildlife Research Unit Technical report no. 4
Title: Spatial distribution of bobcats and gray foxes in eastern Florida
CITATION PAGE IMAGE ZOOMABLE
Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00073791/00001
 Material Information
Title: Spatial distribution of bobcats and gray foxes in eastern Florida
Series Title: Technical report
Physical Description: x, 83 leaves. : ill., maps ; 28 cm.
Language: English
Creator: Labisky, Ronald F
Progulske, Donald Robert, 1956-
U.S. Fish and Wildlife Service
Publisher: Cooperative Fish and Wildlife Research Unit, University of Florida
Place of Publication: Gainesville FL
Publication Date: [1982]
 Subjects
Subject: Bobcat -- Florida   ( lcsh )
Gray fox   ( lcsh )
Wildlife management -- Florida   ( lcsh )
Foxes -- Florida   ( lcsh )
Genre: government publication (state, provincial, terriorial, dependent)   ( marcgt )
bibliography   ( marcgt )
non-fiction   ( marcgt )
 Notes
Bibliography: Includes bibliographical references (leaves 55-59).
Statement of Responsibility: R.F. Labisky, principal investigator ; D.R. Progulske, Jr., project investigator.
General Note: "September 1982."
General Note: "Submitted to: Cooperative Fish and Wildlife Research Unit, University of Florida ; supported by: U.S. Department of the Interior, Fish and Wildlife Service, Contract No. 14-16-0009-79-062."
Funding: This collection includes items related to Florida’s environments, ecosystems, and species. It includes the subcollections of Florida Cooperative Fish and Wildlife Research Unit project documents, the Sea Grant technical series, the Florida Geological Survey series, the Coastal Engineering Department series, the Howard T. Odum Center for Wetland technical reports, and other entities devoted to the study and preservation of Florida's natural resources.
 Record Information
Bibliographic ID: UF00073791
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved, Board of Trustees of the University of Florida
Resource Identifier: aleph - 001891578
oclc - 29860781
notis - AJW6807

Table of Contents
    Title Page
        i
    Acknowledgement
        ii
    Table of Contents
        iii
        iv
    Executive summary
        v
        vi
        vii
        viii
        ix
    Preface
        x
    Spatial distributions of bobcats and gray foxes in Eastern Florida
        Page 1
        Introduction
            Page 1
            Page 2
        Study area
            Page 3
            Page 4
            Page 5
        Methods
            Page 6
            Field collection
                Page 6
                Page 7
            Data analysis
                Page 8
                Page 9
                Page 10
                Page 11
        Results
            Page 12
            Gray foxes
                Page 12
                Page 13
                Page 14
                Page 15
                Page 16
                Page 17
                Page 18
                Page 19
                Page 20
                Page 21
                Page 22
                Page 23
                Page 24
                Page 25
                Page 26
            Bobcats
                Page 27
                Page 28
                Page 29
                Page 30
                Page 31
                Page 32
                Page 33
                Page 34
                Page 35
                Page 36
                Page 37
                Page 38
                Page 39
                Page 40
                Page 41
                Page 42
        Discussion
            Page 43
            Gray fox
                Page 43
                Page 44
                Page 45
            Bobcats
                Page 46
                Page 47
                Page 48
                Page 49
            Gray fox and Bobcat interactions
                Page 50
                Page 51
                Page 52
        Management implications
            Page 53
            Page 54
        Literature cited
            Page 55
            Page 56
            Page 57
            Page 58
            Page 59
    Scent-station indices as indicators of population abundance for bobcats, raccoons, gray foxes, and opossums
        Page 60
        Introduction
            Page 60
            Page 61
        Study area
            Page 62
            Page 63
        Methods
            Page 64
            Page 65
        Results
            Page 66
            Page 67
            Page 68
            Page 69
            Page 70
        Discussion
            Page 71
            Page 72
            Page 73
            Page 74
        Conclusions and recommendations
            Page 75
            Page 76
            Page 77
        Literature cited
            Page 78
            Page 79
            Page 80
    Appendix A: Capture date, sex, age, weight, and measurements for gray foxes...
        Page 81
        Page 82
    Appendix B: Capture date, sex, age, weight, and measurements for bobcats...
        Page 83
Full Text








TECHNICAL REPORT NO. 4


SPATIAL DISTRIBUTION OF BOBCATS

AND GRAY FOXES IN EASTERN FLORIDA



R. F. Labisky, Principal Investigator
D. R. Progulske, Jr., Project Investigator






School of Forest Resources and Conservation
Institute of Food and Agricultural Sciences
University of Florida, Gainesville, FL 32611





Submitted to:



Cooperative Fish and Wildlife Research Unit
University of Florida
Gainesville, FL 32611



Supported by:



U.S. Department of the Interior
Fish and Wildlife Service

Contract No. 14-16-0009-79-062


September 1982




















ACKNCLECGEENIS


This project was funded jointly hy the Florida

Cooperative Fish and Wildlife Research Unit and the School

of Forest Resources and Conservation, University of Florida.

Technical advice was provided ty: Cr. Sicaard W.

Grjeory, Leader, Florida Ccoperative fish and Wildlife

Research Unit; Dr. F. fayne Kiny, Director, Florida State

Iuseum; Dr. Stephen Hiumphrey, Associate Curatcr of

Ecology, Florida State Museum; and James P. Erady,

Supervisory Biologist, bureau of Wildlife Bosearch, Florida

Game and Fresh water Fish Ccimission.

Thanks are also due tne fcllcwinq individuals: Mark C.

Conner assisted with all phases of the research study;

Harold Gcrnto and his starr, University of F~crida researchh

and Education Center at Welaka, expedited field work and

provided excellent facilities; DLuglas Ducant furnished

morai support and occasicual overnight lodqinq; D. W.

Tredinick alicwed access to nis prcperty to radio-tracx

boLcats; and Julie A. Hcvis assisted in tne preparation or

tne report.
















TAELE CF CONIEhIS


PAGE

ACKNOWLEDGMENTS...................................... ii

EXECUTIVE SUMMA3Y....................................... v

PREFACE...................................... ......... x

PART 1
SPATIAL DISTRIBUTIONS CF ECBCATS
AND GRAY FOXES IN EASTERN FLCIL.................... 1

INTRODUCTION..... .............................. 1

STUEY AREA....................................... 3

METHODS ......................................... 6
Field Collection.................... ......... 6
Data Analysis.................................. 8

RESULTS.................... ..................... 12
Gray Foxes......................... ........ 12
Home Range Size................. .......... 12
Core Area Size........................... 1
HUcm Range and Ccre Area Cverlap.......... 14
Activity Patterns......................... 19
Daily Movement Patterns.. ............... 22
Habitat Use. ...... .... ................... 22

Bobcats..................... .............. 27
Hcme Range Size....... ........ ............ 33
Activity Patterns........................... 36
Daily Movement Patterns........ ............ 36
Habitat Use.................. ............. 36

DISCUSSIc ...................................... 43
Gray Fox ........ ............................. 43
Hcme Range Variatility..................... 43
Daily Moveents................................... 45

BoLcats......... ..... ...... .................. 46
H~e Fauige Variability......... ..... .... 46
Daily Hovement ................ ........... 50

Gray Fox and Botcat Interactions............... 50


iii












MANAGEMENT IMPLICATICNS .... ... ............... 53

LITERATURE CITED................................ 55

PAA-T 2
SCENT-STATICN INDICES AS IIDICATCES
OF POPULATION ABUNDANCE FOR
BOBCAIS, RACCOONS, GRAY FOXES, AND OECSSUdS........ 60

INTBODUCTION.................................... 60

STUDY AREA ...................................... 62

METHCDS......................................... 64

RESULTS .............. .......................... 66
Bobcat Visitation..................... ...... 66
Baccoon Visitation ........................... 66
Gray Fox Visitation............................ 69
COossum Visitation............................ 69

DISCUSSION ...................................... 71

CCNCLUSIONS..... ................................ 75

LITELAT7 S CITED............................... 78

APPENDIX A
CAPTURE DATE, SEX, AGE, WEIGHT, AND MEASUEEMENTS
FOR GRAY FOXES CAPTURED ON THE WEIAKA STUDY AREA,
1980-1 82............ ................................. 80

APPENDIX E
CAPTURE LATE, SEX, AGE, WEIGHT, AND MEASUEEMENIS
FOB EOECATS CAPTURE CON THE WELAKA SIUDY AREA,
1980-1 82 ............................... ............ 82
















EXECUTIVE SUMMARY


Part 1: Spatial Distributions of Bobcats and Gray Foxes
in Eastern Florida

Home ranges and movements were determined by

radio-telemetry for a selected populatica or gray foxes

(Drocyon cinereoarienteus) and bobcats (Felis rufus) on a

150-kmz study area in ncrtaeastern ficrida. Six

radio-ccllared gray foxes, Iccated 869 times from December

1980-December 1981, had a mean hcme range size (harmonic

afan analysis) of 582 ha (range=223-1140 ha). The acme

ranges ci male and female rcxes did not differ in size

(?>0.05). Foxes were principally nocturnal; daily movements

dirfered significantly among seasons (P<0.05), hut not

between sexes (P>C.05). Daily movements of cfxes averaged

8242 m during the breeding season (Ceceinber-narch) ; 2235 m

during the denning and helping season (April-July); and

4385 m atter the young had become independent

(Auqust-Ncvember) Simultanecus estimaticn procedures

revealed that tie yray foxes preferred dense areas, such as

pine flatwcods (Pinus elliottii) curing the day, and acved

into more open areas, such as meadows and Icngleaf

pine-turkey oak sandhills (iinus Lalustris-~uercus laevis),

at night. Foxes avoided bcttczland hardwoods totally.

v











Iwc radio-collared male obhoats, located 269 times from

July 1980-December 1981, had a mean home range size

(harmonic mean analysis) of 3820 ha (range=3680-3960 ha).

Although movement activity was recorded during all hours of

the day, bobcats were chiefly nocturnal. Daily movements

averaged 6090 m. Simultaneous estimation procedures

indicated that botcats preferred tottcmland hardwoods during

day and night; a secondary preference for pine ilatwoods was

exhibited during the day. Bobcats generally avoided

habitats witA open understories during the day, and traveled

in habitats with medium understcries at night. No

significant competitive interactions were noted between the

gray foxes and botcats.

It is practical to manage for gray foxes through

habitat manipulation coupled with harvest regulation.

However, due to the solitary tehavicr of bobcats, tne most

efficient way to manage them is tarcuqh population

mcnitoring programs together with manipulation of harvest.











Part 2: Scent-Station Indices as Indicators of
Population Abundance for Eotcats, Raccoons,
Gray Foxes and Cpcssums

Population trends of totcats, raccoons (POcc cn lotor),

gray foxes, and opossums (Didelphis virginiana) were indexed

by scent-station transects on the 918-ha Welaka Reserve.

Three permanent transects were operated for 1 night per

month for the 24-month period, January 1980-December 1981.

Each transect was comprised of 10 stations spaced at 0.32-km

(0.2 mi) intervals; each station consisted of a circle of

sifted sand 0.91 m (3 ft) in diameter with a

centrally-located cottouball that was saturated with totcat

urine. Transects were activated in the afternccn and

checked for visitation the following morning. A visit was

defined as the presence of 1 or more tracks of a species per

station. Mcntaly visitation rates (VB) were calculated for

each species for each transect for each niqLt tuat the

transects were operated. Analysis cf variance procedures

designed to test for differences between years and among

months were applied to arcsine-transfcrmed visitation rates.

Mean monthly scent-station visitation rates for the

24-month period for bobcats, raccoons, gray foxes, and

opossums were 1A, 27., 480, and 1C0, respectively. No

significant differences in the visitaticu rate for bobcats

were detected for years or mcnths (E > 0.05). However,

trends in scent-station visitation rates for bocats did

reflect independently-determined trends Lu pcrulaticn











abundance. Visitation rates ty bobcats were highest in late

fall and winter.

The scent-station visitation rate of 30% for raccoons

in 1980 was significantly ditlerent frcm th 17% rate

recorded in 1981 (P < 0.05). A period of belcw normal

rainfall, beginning in late 1980 and extending throughout

1981, probably concentrated raccoons in wetland habitats

that were not directly indexed by scent-station transects.

Thus, the lower visitation rate for raccoons in 1981, in

contrast to 1980, may have reflected a shift in habitat

utilization rather than a decrease in population abundance.

The highest monthly visitation rate by raccocns occurred in

September.

The scent-station visitation rate oz 521 icr gray foxes

in 1981 was significantly different from the 35W rate in

1980 (P < 0.05). Seasonal trends in gray fox visitation

rates reflected seasonal trends in population abundance,

i.e., rates were highest in fall (Ncvemter), when juveniles

dispersed, and lowest in spring (lay), when the adults fox

population was minimal and young roxes were restricted to

dens.

The erratic trends in monthly scent-station visitation

rates for opposums, as well as the significant interaction

between year and month, suggested that scent-station indices

were not reliable indicators cf trends in opcesum population

abundance.


viii











ITis investigation provided several specific

recommendations for indexing the population abundance of

carnivorous furbearers in Florida by scent-station

transects. If the indices for all species are to be derived

form a single network of transects that is operated once per

year, the network should be operated in October or November

and the transects should be distributed proportionately in

all major habitat types. If species-specific transect

networks are utilized, the network for bobcats and gray

foxes should be operated in Ncvember and the network for

raccoons in September. The large chme ranges or bobcats

during periods of low population density indicated that

scent-staticn indices derived from transects longer than

2.88 km miglt better reflect bobcat population abundance

because mere bobcat ncme ranges wculd be traversed by each

transect. Transects targeted specifically for raccoons

should be distributed so as to sample wetland habitats in

proportion to their availability.















PBEFACE


This report consists of 2 separate and self-contained

papers, which, after appropriate editorial Levisioa, will be

submitted for publication in technical wildlife periodicals.















PART 1: SPATIAL DISTRIEUTICKS CF BCCATS

AND GRAY FOXES IN EASIEBf FICOIDA





INTRCDUCTION





The bobcat (Felis rufus) and the gray fox (Urocyon

ScinereoarSeneus) are important mammalian predators in

Florida. The population status of the tbocat, however, has

become a recent and major concern ct conservationists

because the annual harvest of bobcats in the United States

has risen steadily during the past decade due tc an

increased demand lor pelts cf spotted cats. In Florida, the

average price per bobcat pelt has risen frcm is to $50

(900%) in the past 13 years (Spratt and Brady 1982).

The bobcat is listed on Appendix II of the Convention

on International Trade in Endangered Species of Wild Fauna

and Flora (CITES). This designation requires that countries

exporting pelts must provide information to verify that the

harvest of these animals is not detrimental tc survival of

the species. Recently, a U.S. District Court, using CITES

as a basis, issued an injunction prohibiting the U.S. Fish

and Wildlife Servics tcca authorizing tae eaxcrtaticn of















PART 1: SPATIAL DISTRIEUTICKS CF BCCATS

AND GRAY FOXES IN EASIEBf FICOIDA





INTRCDUCTION





The bobcat (Felis rufus) and the gray fox (Urocyon

ScinereoarSeneus) are important mammalian predators in

Florida. The population status of the tbocat, however, has

become a recent and major concern ct conservationists

because the annual harvest of bobcats in the United States

has risen steadily during the past decade due tc an

increased demand lor pelts cf spotted cats. In Florida, the

average price per bobcat pelt has risen frcm is to $50

(900%) in the past 13 years (Spratt and Brady 1982).

The bobcat is listed on Appendix II of the Convention

on International Trade in Endangered Species of Wild Fauna

and Flora (CITES). This designation requires that countries

exporting pelts must provide information to verify that the

harvest of these animals is not detrimental tc survival of

the species. Recently, a U.S. District Court, using CITES

as a basis, issued an injunction prohibiting the U.S. Fish

and Wildlife Servics tcca authorizing tae eaxcrtaticn of











bobcat pelts. The court indicated it was net satisfied with

the criteria imposed on the states by the U.S. Fisa and

Wildlife Service to determine the suitability cz bobcats for

harvest (Department of the Intericr 1i82).

The general status of the gray fox has not been as

controversial as that of the bobcat because it is more

abundant and is not listed by CIIES. Although harvested in

large numbers throughout the Southeast and in other regions

of the United States, few projects have been devoted to the

study of its biology (Wood et al. 1558, Lord 1961, Trapp and

Hallberg 1975, Root 1581). In Florida, because it is

illegal tc harvest gray foxes and red foxes (Vuljes vuljes),

aucn of the interest in these species has been related to

tLeir role as a carrier of rabies (kccd 1554, Jennings et

al. 1960, Mclean 1970, Carey 1982).

This study was designed to investigate the movement

patterns of bobcats and gray foxes in north Fl1cida via

radio-telemetry; to ascertain the relative use of different

habitats by each species; and to evaluate possible

interspecific competition between the species.















STULY AREA


The Nelaka Study Area, which encompasses approximately

150 kmz, is located in southern Putnam County of

northeastern Florida. The area has little tcpcgraphic

relief, with elevations varying from 3-20 m above sea level.

The soils are principally well-drained sands, although

poorly-drained muck soils cccur in some of the betterr areas

(Laessle 1942, Schultz 1979). Small lakes and po.ds are

common throughout the area. Annual mean temperature and

rainfall for the local region are 22 C and 137 cm,

respectively (National Cceanic and Atmospheric

Administration 1974).

The area is predominantly forested, being comprised of

the following plant communities: hcttcmland hardwoods; pine

flatwoods and pine plantations (Finus elliottii and Pi.us

aliustris); icngleaf pine-turkey oak sandhills (Pinus

palustriis-.uercus laevis) live oak hammocks (Gercus

vZiriniana var. viijiniana) ; orange groves (Citrus spp.);

and open fields. A detailed description of these habitats

is available in Laessle (1942), fMonk (1965), and Veno

(1976).











The ccre study area was tie 9.2 ku2 University of

Florida Welaka Research and Educaticn Center (ceiaka

Reserve), located on the east bank cf the St. Johns River

near the tcwn of Welaka (Fig. 1). The Welaka Reserve is

unique in that approximately 1/3 of the area has been

designated as inviolate since 1939. Inviolate areas have

been protected frcm all man-induced influences except fire

protection and road maintenance. Boundaries of the Reserve

are fenced except along the river. Hunting and trapping

have not been permitted on the Welaka Reserve since the late

1930's.






5











SATSUMA N











CRESCENT
LAKE


SPOMONA PARK


OCALA 8R 309A
NATIONAL 0 |O LK









LITTLE ,
LAKE GEORE Lake
A EO Eet SR 308




2 KM




Fig. 1. Map of the Welaka Study Area, Florida. The
shaded area delineates the 150-km2 Welaka Study Area.
The thin dashed-line represents the boundary of the
Welaka Reserve.















MEIBODS


Field Ccllection

Trapping of gray foxes and bobcats was conducted on the

Welaka Ueserve during January-March 1980, July-December

1980, January-March 1981, and January 1982 (post-study).

Custom-made and commercially available box-type live traps

were used to capture all animals. Twc models ot dead-bait

traps were used, measuring approximately 39 x 39 x 107 cm or

61 x 61 x 122 cm. The live-hait trap dimensicns were 61 x

41 x 122 cm. The principal dead bait was veniscn; live bait

included roosters, guinea fowl, and quail.

Captured animals were simultaneously injected

intramuscularly with an anesthetic, ketamine hydrochloride

(Vetalar--Parke, Davis, and Company, Eetroit, MI) and a

salivation depressant, atropine sulfate (Centaur Company,

Caarlctte, NC). The ketamine hydrochlcride and atropine

sulfate were administered at dosage rates of 11 mj/kg and

0.05 mq/kq body weight, respectively. Induction time was

approximately 4-5 minutes.

immobilized animals were sexed, weighed,. ad measured

(Appendices A and B). Gray foxes and botcats were fitted

with radic-telemetry ccilars (Telcnics, Inc., Yesa, AZ, and















MEIBODS


Field Ccllection

Trapping of gray foxes and bobcats was conducted on the

Welaka Ueserve during January-March 1980, July-December

1980, January-March 1981, and January 1982 (post-study).

Custom-made and commercially available box-type live traps

were used to capture all animals. Twc models ot dead-bait

traps were used, measuring approximately 39 x 39 x 107 cm or

61 x 61 x 122 cm. The live-hait trap dimensicns were 61 x

41 x 122 cm. The principal dead bait was veniscn; live bait

included roosters, guinea fowl, and quail.

Captured animals were simultaneously injected

intramuscularly with an anesthetic, ketamine hydrochloride

(Vetalar--Parke, Davis, and Company, Eetroit, MI) and a

salivation depressant, atropine sulfate (Centaur Company,

Caarlctte, NC). The ketamine hydrochlcride and atropine

sulfate were administered at dosage rates of 11 mj/kg and

0.05 mq/kq body weight, respectively. Induction time was

approximately 4-5 minutes.

immobilized animals were sexed, weighed,. ad measured

(Appendices A and B). Gray foxes and botcats were fitted

with radic-telemetry ccilars (Telcnics, Inc., Yesa, AZ, and











Wildlife Materials, Inc., Carbondale, IL); the transmitter

frequencies were between 150-152 megahertz (MHz), with pulse

rates of 45-60 per minute. The right ear of each animal was

tattooed with an identifying alpha-numeric code. Prior to

release, all anesthetized animals were returned to the traps

until they had regained their equilibrium, which usually

occurred within 1 1/2 hours pcst-injection.

Radio-coliared animals of both species were initially

located at intervals of 4 hours during at least 1 continuous

24-hour period per week. Beginning in August 1981, selected

gray foxes and bobcats were clcated at hourly intervals for

either an 8- or 24-hour period each week. Gray Loxes, which

were principally inactive between 0ECC-1600 hours, could be

monitored for a 24-hour period by clcating them hourly for

8-hour periods between 0000-0800 hours and between 1600-0000

hours. Bobcats, which were mcre active than gray foxes

during the day, were located every hour during the 24-hcur

(OCOO-0COC hours) monitoring periods. Each radio-collared

animal that was not monitored at hourly intervals during a

particular week was located a minimum of 2-3 times that week

to detect any large shifts in its pcsiticn and to verify the

runctioninj of its transmitter.

A trucx-mounted, omni-directional whip antenna was used

to detect the general lccaticn of the radio-ccilared

animals. A 3-element, hand-held yagi antenna vas tnen used

to determine the animal's sypcific location Ly nQans ci











triangulaticn procedures (CcchraL 1580). Radio-location

readings were taken at distances of
and of <0.4 km for bobcats; consistently close approaches

were possible due to the extensive network of reads, trails,

and firelanes. The maximum location error was approximately

1 ha for foxes and 3 ha for bobcats. Each animal location

was plotted on a plastic acetate sheet, overlaid on aerial

photos. Fox and bobcat locations were plotted at photo

scales of 1:7,920 and 1:15,84C, respectively. The date,

time, habitat, weather condition, and animal activity were

recorded for each located animal. Ilctted information was

later converted to x and y coordinates by the use of a

plastic acetate grid-sheet overlaid on the aerial photos.

aadio-lccations were divided into daytime (sunrise to

sunset) and nig4ttime Iccaticns (sunset to sunrise).





Data Analy sis

The home ranges cf gray foxes and bobcats uere

calculated using 3 methcds. Ihe first was the minimum area

method, in whicn a line was drawn connecting the utmost

points of a location distriiuticn (fchr 1I97). The second

method was a modified form of the atypical habitat

elimination technique preferredd Labitat method), in which

the amount cf preferred habitat within each arinal's home

range was calculated (Ailes 1969). ILe third ootLed was the











harmonic mean measure or animal activity areas based on the

harmonic mean of an areal distribution, which used isolines

that were correlated with areas of equal activity (Dixon and

Chapman 1980). The isoline that encompassed 951 of the

radio-locations was considered as the hoce-range boundary.

Isoline intervals were set at 200 m for the gray fox

analysis and at 200 m and 600 m for the bobcat core analysis

and home range analysis, respectively. Open water that

occurred within home ranges was not included in the

calculations of home rage size.

Home ranqe overlap (using the mirimum area and harmonic

mean methods) was measured by calculating 2 values: average

percentage overlap and percentage of radio-locations within

the overlap area. Areas cf greatest activity (care areas)

and centers or activity were determined by utilizing

harmonic mean calculations. The core area boundary was

subjectively defined as the isoline that encompassed >55% c

the radio-locations for a particular individual (Anderscn

1982). The centers of activity were considered equivalent

to the harmonic mean center.

Daily activity patterns were analyzed by grouping the

locations into 2-hour intervals and then deterzizing the

percentage c locations with activity within each 2-hour

interval. lurae distance measures were calculated in

evaluating daily movement patterns: between cest arcas;

forajinj radius; and total daily acvement. Eethen rest











areas was defined as tne distance between tne rest area or

one day and that of the the next aay. Foxes citen departed

and returned to the same rest area. Foraging radius

(calculated only for foxes) was the average distance from

the daytime rest area to each of the subsequent nighttime

locations. Total daily movement was the sum of the

distances between successive locations tarcughout a 24-hour

period. Cnly days for which 6 cr mcre locations per

individual existed were used in tne calculations of the

distance measures.

Gray fox movements were also divided into 3, 4-month

seasons based on information from this and other studies

(Layne and ZcKeon 1956, Sullivan 1956, Wocd 1956, Lord 1961,

Follman 1S73). These seasons were designated as the

breeding season (December-March) ; the denning and helping

season (April-July); and the independence of the young

(August-November). Movement data were analyzed using

analysis or variance procedures (General Linear Models

Procedures-GLM) and Duncan's [ultiple range test from the

Statistical Analysis System (Helwig and Council 1979).

Habitat was divided into 8 overstoiy types and 3

understcry types. 'ie cverstcry types were bcttcmland

hardwoods, live oak harrmocks, live cak scrut (jiyrcus

vir~iniana var. eminata) Icrgleat pine-turkey oak

sandhills, meadows and open areas, mixed wccds, pine

riatwcods, and pine plantaticns (Venc 1976, SchuLtz 1979).











The understory types were open, medium, and dense, as

determined by subjective visual observation. The proportion

of different overstory and understory habitats were

calculated from the aerial-phcto overlays that included all

of the home ranges of the radio-ccllared animals. The

distribution of the radio locations in various habitats was

assumed to reflect the proportionate use of those habitats

by both gray foxes and bobcats. Chi-square analyses were

employed to determine if gray foxes and bobcats used the

various habitats in propcrticn to their availability for the

following categories: total-overstory; day-cverstcry;

night-overstory; total-understory; day-understcry; and

niqnt-understory. If the chi-square tests were significant,

simultaneous estimation procedures, developed by Neu et al.

(1974), were used to determine which habitats were preferred

or avoided. Because simultaneous estimates were performed,

a procatility level of 0.1 was used to prevent individual

ccnzidence interval errors (Neu et al. 1S74).
















RESULTS


Gray Foxs

Fourteen gray foxes (8 males and 6 females) were

captured 29 times in 1,624 trap-nights during the periods

September 1980-March 1981 and January 1982. Three male and

3 female gray foxes were fitted with radic-collars.

The 6 radio-ccllared gray foxes were clcated 869 times

from December 5, 1980-December 31, 1S81. Five cf tae 6

animals accounted tor 865 (>99X) ot the locations (Table 1);

55 (6%) of these locations were ccnfirmed visually. Giv-

hundred thirty-Aive (61E) cf the locations were recorded

during the day and 334 (30i) at night.

A population estimate cf 1.0 yray foxes per kma for tne

winter cf 1391 was derived frcm mark-recapture data

(Schnabel estimate) and radio-telemetry information. The

population probably increased during the summer and fall due

to the addition of ycung animals.





Homq Range Size.--Caiculated home ranges or 5 gray

foxes (2 males and 3 famalts) averaged 400 ha using the

minimum area motacd, 360 "a usinq the ureierred aatitat
















RESULTS


Gray Foxs

Fourteen gray foxes (8 males and 6 females) were

captured 29 times in 1,624 trap-nights during the periods

September 1980-March 1981 and January 1982. Three male and

3 female gray foxes were fitted with radic-collars.

The 6 radio-ccllared gray foxes were clcated 869 times

from December 5, 1980-December 31, 1S81. Five cf tae 6

animals accounted tor 865 (>99X) ot the locations (Table 1);

55 (6%) of these locations were ccnfirmed visually. Giv-

hundred thirty-Aive (61E) cf the locations were recorded

during the day and 334 (30i) at night.

A population estimate cf 1.0 yray foxes per kma for tne

winter cf 1391 was derived frcm mark-recapture data

(Schnabel estimate) and radio-telemetry information. The

population probably increased during the summer and fall due

to the addition of ycung animals.





Homq Range Size.--Caiculated home ranges or 5 gray

foxes (2 males and 3 famalts) averaged 400 ha using the

minimum area motacd, 360 "a usinq the ureierred aatitat
















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method, and 532 ha using tae harmcnic mean method (Tatle 1,

Fig. 2). Ncne of the methods yielded a significant

difference (P>0.05) in home range size between sexes.





Core Area Size.--Gray foxes did not use all portions of

their home ranges with equal frequencyy (Fig. 3). Instead,

their activities were concentrated in certain areas within

taeir hcme ranges (core areas). These core areas probably

contained the most favored resting places, and possibly the

best foraging areas (Jewell 1966, Ever 1968). Sizes of the

core areas ranged from 39-190 ha with a mean of 32 ha.

These areas averaged only 16X of the total hame ranje size

but yielded 57-76% of the radio-locations (Tatil 2).





Hoze -anne and Core Area Cverlar.--CverlaF amcog acme

ranges was variable due to the tendency of gray roxes tc

establish family home ranges (Lord 1961). Degree of overlap

between males and females depended cn their relationship

(dated cr not mated). Mated pairs had the greatest average

area overlap (64, and 60'f) irrespective cf tae acse range

method whereas nou-mated anildis had less area overlap (6/

and 35X) (iabie 3). Tae single male-male ccmbinaticn was

variable defending on the method cf cvorlap calculation (19f

and 567%). female-remsal area over lap was generally lcw (<1

ani 11 ) .





















































Fig. 2. Home ranges of 2 males and 3 female gray foxes,
defined by the harmonic mean method, Welaka Study Area,
Florida, 1980-1981. The harmonic mean center is
represented by the black dot within each home range.
The center is identical for foxes GFI and GM4.


















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Fig. 3. Three-dimensional representation of the home
range of a female gray fox (GF3), illustrating
differential use of areas within the home range
defined by the minimum area method. Each individual
grid square equals 1 ha.
















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Overlap among core areas was variable. One mated pair

(GF1 and GM4) had nearly identical core areas and centers of

activities, whereas the other mated pair (GF3 and GM6) had

widely separated core areas (Iig. 4). This latter

difference occurred because the hcam ranges of GF3 and GM6

also were separate (not a mated pair) until the last 2

months cf the study after which time they were found

together consistently, and considered a mated pair. Slight

overlap of core areas existed for some ncn-mated animals and

for males; however, a review of simultaneous radio-location

data showed that the overlap was spatial and not temporal in

nature. There was no overlap of core areas among females.





Activity Patterns.--Gray foxes were predictaLjy active

at night and sedentary during the day. Activity commenced

shortly before sunset, peaked between 2000-C4CO hours, and

decreased shortly after sunrise. Lhe animals were

essentially sedentary between 110C-1500 hours (Fig. 5).

Activity patterns paralleled the changing periods of

daylight throughout the year. There was no notable

difference between the overall activity patterns at males

and females.


























































Fig. 4. Core areas of 2 male and 3 female gray foxes,
Welaka Study Area, Florida, 1980-1981. The black dot
within each core area represents the harmonic mean
center.


















































































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Daily Movement Patterns.--The daily movement patterns

or gray foxes were repetitive. They traversed a large

portion of their home range each night and often returned to

approximately tie same rest area from whica they departed.

The average distance between rest areas was 426 m; the

average tcraging radius was 779 m; and the average total

daily mcvement was 3179 m (Table 4). There was no

significant difference in daily movements of males and

females (P>0.05).

Daily movements of foxes differed among seasons (Table

5). All movements (between rest areas, foraging radius, and

total daily movement) were lowest during the denning and

wuilping season (April-July) and greatest during the

breeding season (December-March). Ictal daily micvement was

the only variable that was significantly different (P<0.05)

daong all 3 seasons, being lowest during the denying and

helping season (mean=2235 m) and highest during the

breeding season (mean=8242 i).





Habitat Use.--Gray foxes did not use either overstory

or understory habitats in prcFcrticn to their availability

(=<0.01). Moreover, they utilized different habitats during

the daytime and nighttime (Figs. 6 and 7). Ecxes usually

remiainec in areas with d~nse understories, such as pine

flatwoods and live cak scrub, durlrg tae day and moved





















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into more open areas, such as longleaf pine-turkey oak

sandhills and meadows, at night (Figs. 8 and 9).

Simultaneous estimation procedures (Neu et al. 1974)

supported the initial findings that fcxes preferred dense

habitats, suca as pine flatwccds and live oak scrub, during

the daytime (P<0.1), and open habitats, such as meadows and

lougleaf pine-turkey oak sandhills, at night (P<0.1)(Tables

6 and 7). Furthermore, irrespective of the time of day,

foxes avoided bottomland hardwoods and live cak aammccks and

preferred live oak scrub. Cther habitats were either

avoided or showed non-significant use.





Bobcats

Seven bobcats (4 males and 3 females) were captured 21

times in 2,657 trap nights. Six animals (3 maies and 3

tamales) erre fitted with radio-cellars.

The 6 collared bobcats were located 294 times from

February 15, 1980-Decemfer 31, 1981. Two males (BM2 and

314) accounted for 269 (925) of the bobcit radic-locations.

Three of tie animals radio-ccllared during January-June,

1980 (Table 8), were lost from the study due tc

radio-ialtunctions or deaths. Cne hundred seventy-nine

radio-locations (b1%) were recorded during the day and 115

(39.) at nijiht.
































60







PFf 50
40


20




Fig. 8. Daytime habitat use by a female gray fox (GFl),
Welaka Study Area, Florida, 1980-1981. Habitats are
pine flatwoods with a dense understory (PFLT/DNS) and
longleaf pine-turkey oak sandhills with an open understory
(LPTO/OPN). The broken black line is the boundary between
the habitats. The arrows point to extremes in location
frequencies. One grid square equals 1 ha.































30 p
050






40
% s o


20



Fig. 9. Nighttime habitat use of female gray fox
(GF1), Welaka Study Area, Florida, 1980-1981.
(See Fig. 8 for explanation of symbols.)















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A population estimate cf 0.52 totcats per kma was

calculated from trapping, radio-isotope tagging, and

radio-telemetry data before an apparent die-cft probablyy

due to disease) occurred in June 1960, after which time the

population density decreased (Ccnner 1982).





Homeg angel Size.--Home ranges of the 2 male bobcats

averaged 4438 ha using the minimum area method, 3350 ha

using the preferred habitat method, and 3820 ha using the

harmonic mean method (Fig. 10). These 2 animals had an

average hcme range overlap of 101 and 8%, based oa the

minimum area and harmonic mean methods, respectively (Table

8).

Bobcats, like the gray ioxes, visited different

pcrticns of their acme ranges with varying r requency (Fig.

11). Their core areas, however, were act as distinct as

those of tne gray foxes. Tae mean core area ct the 2 male

individuals was 909 aa (range=695-1124 ha), whicn accounted

for an average of 24 ct their hcme range size and an

average or 59% the total radic-locations. No overlap was

determined icr core areas between ancials (Fig. 10).























































Fig. 10. Home ranges and core areas of 2 male bobcats,
defined by the harmonic mean method, Welaka Study Area,
Florida, 1980-1981. The core areas are delineated by
the dotted lines within each home range. The harmonic
mean center is represented by the black dot within each
home range.





















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25
0
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Fig. 11. Three-dimensional representation of the home
range of a male bobcat (BM4), illustrating differential
use of areas within the home range. The bold black
line is the home range boundary defined by the minimum
area method. Each individual grid square equals 4 ha.











Activity Patterns.--Most botcat activity occurred at

nig.t; however, bobcat nighttime activity was more erratic

than that ot the gray fox. Activity began to increase

before sunset, with peaks between 18GC-2000 and 2200-0000

hours, decreased to a nighttime low between 0200-0400 hours,

and then increased about sunrise. Eocat activity was

recorded during all periods of the 24-hour day (Fig. 12).





Daily Movement Patterns.--The daily movement patterns

or the tobcat were nomadic within their hcme range;

movements of varying distances occurred at any time ot the

day or night. The 2 male ottcats had a mean distance

between rest areas ot 2700 w (range=0-4245 m, SE=543 m) with

an average total daily mcvement ot 6090 m (range=412-9631 m,

SE=1027 m).





Habitat Use.--Bobcats did not use either overstory or

understory habitats in propcrtica tc their availability

(0<0.01). They used habitats with dense or medium

unaerstcries and avoided naaitats witA cpen understories.

Moreover, tae bobcats utilized different habitats during the

day than at night, although these differences were not as

distinct as those of the gray fox (Fiqs. 13 and 14).












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Simultaneous estimation procedures (Neu et al. 1974)

indicated that bobcats preferred hottcaland hardwoods hcth

day and night, with some preference toward pine flatvcods

during the day (P<0.1) ablee 9). Ihey avoided areas with

open understories, such as meadows and longleaf pine-turkey

oak sandhills, during the day, and traveled in habitats with

medium understories at night (P<0.1) (lable 10). TMeir use

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DISCUSSION


Gral Fox

Home Range Variability.--The mean home ranges of gray

foxes reported in this study (males=602 ha, females=365 ha)

were substantially larger than these reported from other

regions of the United States, except for Misscuri (676 ha)

(Table 11). Although differences in sizes cf home ranges

may be attributed to a number of ractcrs, differences among

habitats, which affect prey distribution and abundance, and

i: turn, rox density (Wood et al. 1958, Trani 1980),

probaiby constitute the singlemcst important variable in

detrmininiq hcme range size cr gray fcxes. The gray fox is

omnivorous; its annual diet is ccaprised primarily of

vertebrates (50) vegetaticn--imotly fruit (15A), and

arthropcds (35s) (Yoho and Henry 1972, Trapp 1973, Trani

1980). Open habitats provide areas in which gray foxes

forage for food. Habitats with dense understcries are

important as daytime rest areas but are poor reservoirs of

food supplies. Home ranges of gray cfxes were larger where

habitats were more homogeneous (Harcldscn 1982, this study)

than where taey were more disjunct and interspersed (Trapp

1973, Hallerg 1974, Fuiler 1978, Yearsley and Samuel 198C).















DISCUSSION


Gral Fox

Home Range Variability.--The mean home ranges of gray

foxes reported in this study (males=602 ha, females=365 ha)

were substantially larger than these reported from other

regions of the United States, except for Misscuri (676 ha)

(Table 11). Although differences in sizes cf home ranges

may be attributed to a number of ractcrs, differences among

habitats, which affect prey distribution and abundance, and

i: turn, rox density (Wood et al. 1958, Trani 1980),

probaiby constitute the singlemcst important variable in

detrmininiq hcme range size cr gray fcxes. The gray fox is

omnivorous; its annual diet is ccaprised primarily of

vertebrates (50) vegetaticn--imotly fruit (15A), and

arthropcds (35s) (Yoho and Henry 1972, Trapp 1973, Trani

1980). Open habitats provide areas in which gray foxes

forage for food. Habitats with dense understcries are

important as daytime rest areas but are poor reservoirs of

food supplies. Home ranges of gray cfxes were larger where

habitats were more homogeneous (Harcldscn 1982, this study)

than where taey were more disjunct and interspersed (Trapp

1973, Hallerg 1974, Fuiler 1978, Yearsley and Samuel 198C).









44













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This relationship between hcme range size and habitat

heterogeneity was apparent also cn the Welaka Study Area.

Home ranges containing tae smallest proportion cf

homogeneous dense forested areas (GF1, GF4, GM4) were

smallest (136-293 ha), whereas these with the greatest

proportion of uniform dense crested areas (G53 and Gi6)

were largest (667 and 917 ha). Therreore, cptiaum fcx

habitat is characterized by an interspersicn ct dense

habitats for daytime resting and cpen areas for nighttime

foraging. Intuitively, the proportion cf foraging habitat

should invariably exceed that of resting habitat.





Daily joveentjs.--Foxes were principally nccturnal but

exhibited some crepuscular and early morning activity.

Their activity during the twilight and early daytime hours

may add an important feature to their niche hy allowing their

to oxplcit diurnally-active prey species, such as lizards

and certain small mammals (Trapp 1973). Daily movement

patterns were similar for males and females Lut varied

seasonally. Movement was greatest during the breeding

season. TIese Aindings were similar to these ct Trapp

(1973) in Utah, and Fcilman (1973) in Illiicis, sno reported

that gray foxes moved greater distances in winter (mating)

than summer (denning). Restricted movements Ly foxes in

summer are attributable tc the dependence cf young and the











ready availability cf food (fruits and insects). :n winter,

juveniles are dispersing frox their natal home ranges,

adults may be moving to find mates, ana food may be more

difficult to obtain.





Bobcats

Home Range Variability.--The reported mean size of

bobcat home ranges has varied considerably, from 97 ha tor

female bobcats in Louisiana (Hall and Newsom 1976) to 620C

ha for male bobcats in northern Minnesota (Eerg 1981) (Table

12). Male chme ranges are consistently 2 tc 3 times larger

taan thcse of females. Although sales have identifiablE

core areas, they travel more throughout their hCme ranges

than females (Kitchings and Story 1981).

Home ranges are consistently larger in the northern and

western portions of tne country than in the South (Table

12). This geographical variation in home range size may be

a function of the climate, which, in turn, affects the

variety and abundance of prey. Bobcats are exclusively

carnivorous, leading primarily on mammalian prey (Progulske

1955, Buttrey 1974, Fritts and Seaiander 1978, Guenther

1580). Ihe warn southern regions probably nave a more

consistent year-round prey base thaL the more northern and

western sections of the country.

















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The average home range of the bobcats monitored in this

study was much larger than tncse previously reported from

the southern United States, except for lennessee (Kitchings

and Story 1981). These researchers maintained that the

difference between their study area and the other southern

study sites was probably the richness of the prey base,

waich was related to the prevalence of early successional

fields (old fields). Buie et al. (1981) noted that the

decrease in old field sites may have caused the expansion of

bobcat hcme ranges cn the Savannah Fiver Plant during the

past 20 years. In this study, the reasons for large home

ranges may be two-fcld. First, the number of small mammals

may be relatively low due to the dominance cf homogeneous

dense forested habitats. Seccndly, the apparent die-cfi of

the resident bobcats during the summer of 1980 may have

created vacant habitat into which surviving bctcats expanded

their home ranges. Initial data, based cn trapping and

limited radio-telemetry, showed that the original resident

female bobcats on the Welaka Reserve occupied smaller home

ranges (2CO-1000 ha) than those determined tor the 2 males

(B32 and B34) carter the die-off. Similar range expansions

following bobcat deaths have been reported by Miller (1980)

and LeAbeck and Gculd (1981). Therefcre, qecgrapnicai

variation in home range size of boccats is probably a

function of both habitat and climate, which, in turn affects

the prey base; however, Iccalized factors, such as disease











and exploitation pressure also play a role in affecting home

range size.





Daily Movements.-- Bobcats were principally nocturnal

but were found to be active at any time during the day.

Occasionally they were active all night, but often they had

movement lulls between 00CC-C400 hours. This pattern of

activity is consistent with that reported for bbccats in

south Florida (Guenther 1980) and in Alabama (Miller 1980).

The bobcat activity cycle usually matches that of its prey,

such as cottontail rabbits (Sylvilaqus iloridanus) (Lord

1964) and deer (Odocoileus viriLnianus) (Halls 1978).





Gral Fox and Ectcat Interacticns

The canids and the telids have had a lcng, separate

evolutionary history (Kleiman and Eisenbery 1973). The

feeding and hunting behavior of the canids (omnivorous and

prey chasing) has encouraged a social existence. The basic

social structure consists of a long-lived pair tond in which

the male becomes involved in the rearing of the young. The

pair bend of the gray fcx is year-rcund.







51



The feeding and hunting behavior cf most felids

(exclusively carnivorous with ambush attack of their prey)

has encouraged the development of a social system consisting

of nearly exclusive home ranges (especially between like

sexes) and a solitary existence (Kleiman and Eisenberg

1973). The pair bond of the bobcat generally lasts only

through the breeding season, with the responsibility of

raising the young falling entirely to the female (Bailey

1974).

In this study, differences in gray rcx and bobcat

movements and behavior may have prevented competition

between the species. The nome ranges of male bobcats (3820

ha) were 6-7 times larger than those cf gray foxes (582 ha).

Furthermore, these 2 species had ncticeatle differences in

habitat selection. Bobcats preferred dense bcttomland

hardwood habitats during all portions of the day and night,

and generally avoided open wooded areas, suca as longleaf

pine-turkey oak sandhills. Gray foxes, however, avoided

bottomland hardwood habitats and preferred cpen areas such

as longleaf pine-turkey oak sandhills at night. The single

habitat that was preferred by both species was the dense

pine fiatwcods which was prohbaly used as daytime retreats.

However, due to the prevalence of this type of habitat on

tae study area, it is unlikely that competiticL for pine

flatwcod sites occurred. It is cpssible that the bobcats

precluded the gray toxes iron the bcttomland sites; however,











on the Welaka Reserve where the bobcats visited only

sporadically, gral roxes still avoided bottcmland hardwoods.

Foraging Vabits cf bobcats and gray foxes are similar

during certain periods of the year. Although gray foxes

feed heavily on fruits and insects in summer, they prey

substantially on small mammals, which are also prey of

bobcats, in winter. Habitat selection by the 2 species,

however, may mitigate the potential foraging competition.

Bobcats prefer bottomland sites, which are richer in prey

than forested upland areas (Kitchings and Story 1981, M. C.

Conner and D. R. Progulske, Jr., unpublished data), whereas

gray foxes avoid bottomland habitats, preferring, instead,

upland sites.

In conclusion, although interactions between gray foxes

and bobcats may have occurred, differences in hcme range

size, habitat selection, and foraging behavior probably

prevented significant competition between these 2 species on

the Welaka Study Area.
















MANAGEMENT IMPLICATICNS


Differences in social structure and movements of gray

foxes and bobcats place the species at opposite ends of the

management spectrum. Although both species can be managed

through adjustments in harvest'regulations, it is feasible

to manage gray foxes, but probably not bobcats through

habitat manipulation.

Gray foxes are short-lived omnivcres with definable

naoitat requirements and relatively small hcme ranges.

Consequently, it is possible tc manage for this species

through habitat manipulation coupled with harvest

regulation. Optimum fox habitat has an interspersion of

dense areas for daytime retreats and open areas for

nighttime fcraqing. These habitat tracts shcuid be mixed in

relatively small patches ci 5-15 ha (Lased on a mean home

range size of 300-500 ha) to support the greatest numbers of

animals.

Bobcats are solitary, wide-ranging, and Icng-lived

carnivores. Although habitat preferences cf bbccats can be

identified, tae management of habitats cn the large tracts

of land needed to achieve optixua population levels would

be, in most instances, financially and Icgistically







54



impractical. The most etticient way to manage botcat

populations is probably through pcEulaticn monitoring

programs, such as scent-statica surveys, coupled with the

manipulation of harvest. However, it a decision were made

to manage habitats for bobcats, the preservation oc

bottomland hardwoods would be a critically important

strategy.















LITEiATURE CIiTE


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vulpes). J. mammal. 50:1C8-12C.

Anderson, D. J. 1982. The home range: a new ncnparametric
estimation technique. Ecology. 63:103-112.

Bailey, I. N. 1974. Social organization in a bchcat
population. J. iildl. Manage. 3E:435-466.

Berg, W. E. 1981. Ecology of bobcats in northern Minnesota.
Pages 55-61 in Bobcat research conference proceedings:
current research on biology and management of Ljnx
rufus. Natl. Will. Fed. Sci. and Tech. Ser. 6.

Buie, D. E., I. T. Fendley, and H. F cNab. 1981. Fall and
winter home ranges of adult bobcats on the Savannah
River Plant, South Carolina. Pages 42-46 in Bobcat
research conference proceedings: current research on
biclogy and management to Lynx rutus. Natl. Will. Fed.
Sci. and Tech. Ser. 6.

Buttrey, G. W. 1974. Fcod habits and distribution of tne
bobcat, Lynx rufus rutus (Schreber), on the Catoosa
wildlife Manaqement Area. M.S. Thesis. Tennessee Tech.
Univ., Cookeville. 64 Fp.

Carey, A. B. 1982. The ecology of red foxes, gray ioxes, and
rabies in the eastern United States. Wild. Soc. Bull.
10: 1-26.

Cochran, W. W. 1980. Wildlife telemetry. Pages 507-520 in
S. S. Schemnitz, ed. wildlife management techniques
manual, 4th ed. The Wildl. Scc., Washingtcn, D.C.

Conner, M. C. 1982. Determination of bobcat (1ynx rutus)
and raccoou (rocycn Ictor) population abundance by
radio-isotope tagging. M. S. Thesis. Univ. of Florida,
Gainesville. 55 pp.

Crowe, E. M. 1972. The presence of annuli in bobcat tooth
cementum layers. J. Aildl. Manage. 36:1330-1332.

1975. Aspects oa ageing, growth, and reproduction cf
bobcats froa Wyoming. J. Mammal. 56:177-198.











Department of the Interior. 1982. Expcrt of bobcat taken in
tne 1981-1982 season. Fed. Reqist. 47:1294.

Dixon, K. R., and J. 2. Chapman. 1980. Harmonic mean measure
of animal activity areas. Ecclcgy 61:1040-1044.

Ewer, R. F. 1968. Ethology of mammals. Plenum Eress, New
York, L.Y. 418 pp.

Follman, E. H. 1973. Comparative ecology and behavior of red
and gray foxes. Ph.D. Thesis. Southern Illinois Univ.,
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Fritts, S. H., and J. A. Sealander. 1978. Diets of bobcats
in Arkansas with special reference to age and sex
differences. J. Wildly. Manage. 42:533-539.

Fuller, T. D. 1978. Variable hcme-range sizes o female gray
foxes. J. Mammal. 59:446-449.

Guenther, D. D. 1980. Home range, social organization, and
movement patterns of the bctcat, _Lynx rufus, from
spring to fall in south-central Florida. M.S. Thesis.
Univ. South Florida, Tampa. 66 pp.

Hall, H. T., and J. D. Neusom. 1976. Summer hcme ranges and
movements of bobcats in hottcaland hardwoods of
southern Louisiana. Proc. Southeast. Assoc. Game and
Fish Comm. 30:427-436.

laillberg, D. L. 1974. A contribution toward the better
understanding of gray fox (Uro1cn cinereoargenteus)
temporal and spatial natural history. M.S. Thesis.
California. State Univ., Sacramentc. 280 pp.

Halls, I. K. 1978. White-tailed deer. Pages 43-65 in J. L.
Schmidt and D. L. Gilbert, eds. fig game or North
America. Stackpole Bcoks, Harristurg, Pa.

Haroldscn, K. J. 1982. Habitat ecology of the gray fox in
the czark highland. M.S. Thesis. Univ. Missouri,
Columbia. 99 pp.

Helwig, J. I., and K. A. Council. 1979. SAS users' guide.
1979 edition. SAS Inst. Inc., Cary, N.C. 494 pp.











Jennings, 1. L., N. J. Schneider, A. I. Lewis, and J. E.
Scatterday. 1960. Fox rabies in Flcrida. J. Wildl.
Manage. 24:171-179.

Jewell, P. A. 1966. The concept of home range in mammals.
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Kitchings, J. T., and J. E. Story. 1981. Home range and diet
of bobcats in eastern Tennessee. Pages 47-52 in Bobcat
research conference proceedings: current research on
biolgcqy and management of Lynx rufus. Natl. Widl. Fed.
Sci. and Tech. Ser. 6.

Kleiman, D. G., and J. F. Eisenberg. 1973. Comparison of
canid and telid social systems from an evolutionary
perspective. Anim. Behav. 21:637-659.

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area. Univ. Florida Bicl. Sci. Ser. 4:1-143.

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rox and gray fox reproduction in New York. N.Y. Fish
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Lembeck, M., and G. I. Gould. 1981. Dynamics of harvested
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Lord, B. D., Jr. 1961. A population study of the gray rcx.
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194. Seasonal changes in the activity of penned
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ranges of bobcats as determined ty radio-tracking in
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20:206-214.

McLean, R. G. 1970. Wildlife rabies in the United States:
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Miller, S. D. 1980. The ecolcgy of the bobcat in south
Alatama. Ph.D. Thesis. Auburn Univ., Auburn, Ala. 156
pp.











Monr, C. C. 1947. Table or equivalent populations of North
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Winstcn, Oreg.

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Mirec.

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1980-1981. Florida Game and Freshwater Fish Comm. 7
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Sullivan, E. G. 1956. Gray tox reproduction, denning, range,
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southwest Utah carnivores: Bassariscus astutus and
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Wisconsin, Madison. 251 pp.











_...., and D. L. Hallberg. 1975. Ecclcqy of the gray fox
(Urocyon cinereoarjenteus): a review. Pages 164-178 in
M.W. Fox, ed. The wild canids. Van Nostrand Reinhold
Co., New York, N.Y.

Vano, P. A. 1976. Successional relationships of five Florida
plant communities. Ecology 57:498-508.


Wood, J. E. 1954. Investatgation c fox populations and
sylvatic rabies in the Scutheast. Trans. North Am.
Wildl. Conf. 19:131-139.


1958. Age structure and productivity of a gray fox
population. J. Sammal. 39:74-86.

__, D. E. Davis, and E. V. Komarek. 1958. The distribution
of fox populations in relation to vegetation in
southern Georgia. Ecology 39:160-162.

Yearsley, E. F., and D. E. Samuel. 1980. Use of reclaimed
surface mines by fcxes in West Virginia. J. Wildly.
Manage. 44:729-734.

Yoho, N., and V. Henry. 1972. Foods o! the gray fox (Utrocyon
cingreoarqenteus) on European wild hog (Sus scrofa)
range in east Tennessee. J. Tennessee Acad. Sci.
47:77-78.

Zezulak, D. S., and R. G. Schwab. 1981. A comparison of
density, home range and habitat utilization of bobcat
populations at Lava Beds and Jcshua Tree National
Monuments, California. Pages 74-79 in Ectcat research
conference proceedings: current research on biology and
management of Lynx rufus. Natl. Wildl. Fed. Sci. and
lech. Ser. 6.















PART 2: SCENT-STAIION INDICES AS INDICAIOBS

OF POPULATION ABUNDANCE FCO

BCBCATS, RACCOONS, GRAY FCXES, AND CPCSSUMS





INTRCDUCTION





The use Of scent-station transects as a means of

determining seasonal and annual trends in the relative

abundance of mammalian carnivores has escalated in recent

years. Originally developed as a method for determining the

relative abundance of red (Vulpes vules) and gray foxes

(Uroc2on cinereoargenteus) (ichards and Hine 1953, Wood

1959), the technique has been applied to coyotes (Canis

latrans) (Linhart ana Knowltcn 1975, Davison 1981, Morrisca

et al. 1981), bobcats (Felis rufus) (Brady 1981, Hon 1981,

Johnson 1981, Knowlton and Tzilkowski 1981, Sumner and Bill

1980, Morrison et al. 1981), wolves (Canis lupus) (Pimlctt

et al. 1S69), and river otter (Lutra canadensis) and mink

(Mustela visou) (Humphrey and Zinn 192). Research has

focused also on standardizing and improving otth the

technique and analysis of data (Rcughtcn and Sweeny 1982).















PART 2: SCENT-STAIION INDICES AS INDICAIOBS

OF POPULATION ABUNDANCE FCO

BCBCATS, RACCOONS, GRAY FCXES, AND CPCSSUMS





INTRCDUCTION





The use Of scent-station transects as a means of

determining seasonal and annual trends in the relative

abundance of mammalian carnivores has escalated in recent

years. Originally developed as a method for determining the

relative abundance of red (Vulpes vules) and gray foxes

(Uroc2on cinereoargenteus) (ichards and Hine 1953, Wood

1959), the technique has been applied to coyotes (Canis

latrans) (Linhart ana Knowltcn 1975, Davison 1981, Morrisca

et al. 1981), bobcats (Felis rufus) (Brady 1981, Hon 1981,

Johnson 1981, Knowlton and Tzilkowski 1981, Sumner and Bill

1980, Morrison et al. 1981), wolves (Canis lupus) (Pimlctt

et al. 1S69), and river otter (Lutra canadensis) and mink

(Mustela visou) (Humphrey and Zinn 192). Research has

focused also on standardizing and improving otth the

technique and analysis of data (Rcughtcn and Sweeny 1982).











Despite widespread use of the technique (Jchnson et al.

1981), the relationship between pcpulaticn abundance and

scent-staticn indices remains unclear. To date, tae only

reported correlation between scent-station indices and

population abundance was developed for coyotes in Utah and

Idaho (Eavison 1981).

The objectives of this project were: (1) to evaluate

scent-staticn indices as indicators of seasonal and annual

trends in the abundance of bobcats, gray foxes, raccoons

(Procyon lotor) and opossums (Didelphis virginiana); and

(2) to compare tce indices with population abundance

estimates based on trapping, radioisotope tagging, and

radio-telemetry.















STUEY AEEA


All scent-station data were collected on the University

of Florida Besearch and Education Center at Welaka, Florida.

The 918-ha area is located on the east bank of the St. Johns

River in southeastern Putnam County. Annual mean

temperature and rainfall for the northeastern Florida area

are 22 C and 137 cm, respectively (Eational Cceanic and

Atmospheric Administration 1974).

The soils and plant communities of the Welaka Study Area

were described in detail by Laessle (1942), Veno (1976), and

Schultz (1979). The thick well-drained to moderately

well-drained acid sands were derived from marine sediments.

The area has little topographic variation; elevations range

from sea level in the western sector (St. Johns River) to

20 m in the eastern sector. The increasing relief, and

accompanying decrease in soil moisture, along the

west-to-east gradient gives rise to 3 major plant

communities. Bottomland hardwocds occupy 188 ha (20%) of

the study area and are found principally in the St. Johns

Eiver floodplain; pine flatwccds, primarily

Pinus elliottii var. elliottii, occupy 489 ha (531) and are

located adjacent to the floodplain forest; longleaf pine











(Pinus Lalustris)-turkey oak (uercus laevis) sandhills

occupy 115 ha (13;) and lie in the undulating eastern

portion of the area, adjacent to the pine flatwcods. In

addition, numerous marsh-like ponds, ihich occupy a total of

about 60 ha (6%) of the area, are distributed throughout the

pine flatwoods and sandhills. The remaining 8 of the area

is contained in wcrk and research facilities.

Hunting and commercial trapping have been prohibited on

the Welaka Area since the late 1930s.















METHODS


Three permanent scent-station transects were established

along trails on the Welaka Area; 2 transects were located in

pine flatwoods and 1 in longleaf fine-turkey oak sandhills.

Each transect consisted of 10 stations spaced at 0.32 km

(0.2 mi) intervals; each station consisted of a circle of

moist sifted sand 0.91 m (3 ft) in diameter with a

centrally-placed cottonuall that was saturated with bobcat

urine (Cronk's Outdoor Supplies, Wiscasset, ME). Transects

were operated for 1 night per month for the 24 months,

January 1980-December 1981. Transects were activated in the

afternoon and checked for visitation the following morning.

If rain rendered the traasects inoperative, the procedure

was repeated until an operative transect-night was achieved.

A visit was defined as the presence of 1 or more tracks

of a species per station. Visitation rates were expressed

as the percentage of stations visited by each species for

each transect night.

Analysis of variance procedures were executed cn arcsine

transformed visitation rates using the General linear Models

Procedure (GLa) of the Statistical Analysis System (SAS).

The effects of year and month were tested; the effects of











habitat could aot be tested because there was no replication

of transects in tne sandhill habitat. All statistical

analyses were performed on the transfcrmed rates; however,

in this paper, only the actual percentage visitation rates

are presented.

Population abundance estimates fcr bobcats and raccoons,

with which the scent-station indices were compared, were

derived principally from a companion study (Conner 1982).

Basically, the abundance estimates were based on the

radioisctcpe mark-and-recapture technique (bobcats and

raccoons) and traditional mark-and-recapture methodologies

based on trapping (bobcats, raccoons, gray foxes, and

opossums). Population abundance data for bobcats and gray

foxes were supplemented by radio-telemetry studies

(Proqulske 1982).















RESULTS


Bobcat Visitation

The mean monthly scent-station visitation rate for

bobcats for the 24-months was 1%. Visitaticn by bobcats

never exceeded 4% for any single month (Fig. 1). No

significant trends in bobcat visitation by year or month

were observed (P > 0.05, Table 1).



Raccoon Visitation

The mean monthly rate of scent-station visitation by

raccoons for the 24-months was 271. The visitation rate by

raccoons differed significantly between years (P < 0.01,

Table 1), being greater in 1980 (30j) than in 1981 (17%)

(Fig. 1). The combined monthly rates for the 2 years

indicated that the highest visitation rate was recorded in

September (41%), and tae lowest in Novemter (10c) (Table 2).












'BOBCAT 1980
1981 ---




,----' '---------' \


* -


I
/
\ !


"GRAY FOX A


80.0


60.0


40.0 -


20.0


0.0


I'
/ \ /


'OPOSSUM



A /


JAN MAR MAY JUL SEP NOV


MONTH


Fig. 1. Monthly scent-station visitation rates for bobcats,
raccoons, gray foxes, and opossums, Welaka Area, Florida,
1980-1981.


0.0

40.0


20.0


/ \
* \


/-/ '


2.0











Table 1. Results of analysis of variance Frocedures for arcsine
transformed scent-station visitation rates for hcfcats, raccoons,
gray foxes, and opossums, Welaka Area, Florida, 1980-1981.


Species Factor df F-value



Bobcat Year 1 0.13
Month 11 1.09
Year*Month 11 0.45

Raccoon Year 1 10.33 a/
Month 11 1.78
Year*Mcnth 11 1.S8

Gray fox Year 1 6.53 b/
Month 11 1.84
Year*Month 11 0.26

Cpossum Year 1 8.42 a/
Month 11 1.75
Year*Month 11 2.61 b/


a/P < 0.01.

b/P < 0.05.













Gray Fox Visitaticn

The mean monthly scent-station visitation rate by gray

foxes fcr the 24-months was 48%. The visitation rate by

foxes differed significantly between years (P < 0.05,

Table 1), being higher in 1981 (527) than in 1980 (35%).

The combined monthly rates for the 2 years indicated that

the highest visitation rate occurred in November (82%), and

the lowest rate in May (22%) (Table 2).



CEossum Visitation

The mean monthly scent-station visitation rate by

opossums for the 24-months was 9.9%. Although the

visitation rates by opossums differed significantly by years

(P < 0.01, Table 1), being greater in 1980 (12.5%) than in

1981 (7.3%) (Fig. 1), the significant interaction between

year and month (jP < 0.05, Table 1) nct only negated the

year-difference but alsc further statistical comparisons.











Table 2. Results of Duncan's Multiple Range Test for arcsine
transformed scent-station visitation rates by month for
raccoons and gray foxes on the Welaka Area, Florida,
1980-1981.



Raccoon Gray Fox

Month Rate (%) a/ Month Rate (%) a/

Sep 41 A Nov 82 A
Oct 33 AB Dec 70 AB
Jun 33 AB Cot 67 AB
May 31 ABC Jan 46 ABC
Feb 27 ABC Jun 44 ABC
Aug 21 ABC Feb 42 ABC
Apr 21 ABC Sep 40 ABC
Dec 19 ABC Aug 39 ABC
Mar 18 ABC Mar 35 BC
Jul 17 ABC Jul 35 BC
Jan 15 BC Apr 28 BC
Nov 10 C May 22 C



a/ The presented visitation rates (%) are actual mean
monthly rates, not arcsine transformed rates; those
rates followed by the same letter are not
significantly different f( > 0.05)-















DISCUSSION


Determination of the relationship between scent-station

indices and population abundance is an essential step in

determining the sensitivity of the indices to changes in

abundance. Scent-station indices can detect area-wide

changes in abundance, changes in habitat use, and seasonal

trends in population abundance.

Although the monthly scent-station visitation rate for

bobcats was low throughout the 24-mcnth period (Fig. 1),

trends in the visitation rate were substantiated by

estimates of population abundance and by scat-count indices.

During the 5-month period, January-May 1980, the population

density of bobcats on the study area was estimated at 5

animals (0.52 per km2), the scat-count index was 0.17 scats

per km of trail (Conner 1982), and the scent-station

visitation rate was 1.3%. During the following 16 months,

June 190E-September 1981, the density of bobcats was

substantially lower. In June 1980, 3 of the 5 bobcats on

the area died or disappeared, and only 2 animals were known

to have visited the area periodically from June

1980-September 1981 (Progulske 19S2). Concurrently, the

scent-station visitation rates and scat-count index were











markedly lower, 0.44X and 0.02 scats per km, respectively.

During the 3-month period, Cctober-recember 1981, the bobcat

population density increased as indicated by frequent

observations of unmarked animals and by the capture of an

unmarked male. Similarly, the scent-station visitation rate

and scat count index rose to their highest levels, 1.5% and

0.20 scats per ks, respectively. Thus, both increases and

decreases in bobcat population abundance were reflected by

scent-station indices.

The population density of racccns on the study site was

estimated at 95-105 animals (10.3-11.4 per kmz) during the

winter period, January-March 1981 (Conner 1982). The rate

of scent-station visitation for tais population was 14%

during the same time interval. This 14% visitation rate,

however, was substantially less than the 32% rate for the

January-March period of 1980. The difference in rates

between years may have resulted from rainfall patterns;

below normal rainfall occurred throughout the 2nd half of

1980 and all of 1981. Consequently, in 1981, wetland

habitats cn the study area were diminished in tctal area and

restricted in distribution to the bcttomland hardwoods and

upland ponds, neither of which were mcnitored directly by

scent-staticn transects. The combined effects of the

disproportionate use by raccocas of wetland over upland

habitats on the Welaka Area (Conner 1982), the limited

availability of wetlands in 1981, and the skewed upland











distribution of scent-station transects resulted in

decreased scent-station visitation by raccoons from 1980 to

1981. Sumner and Hill (198C) also noted decreased

visitation of upland scent-station transects by raccoons

when the aniamls were seasonally concentrated in wetland

habitats. Thus, scent-station transects appeared to not

only reflect changes in raccoon abundance, but also shifts

in habitat use.

The population density of gray foxes on the Welaka Area

and surrounding lands was estimated at 1 per km2 during the

winter period, December 1980-March 1981 (Progulske 1982).

Tae corresponding scent-staticn visitation rate for this

breeding population was 56%. During the next 2 seasons,

April-July and August-November 1981, scent-station

visitation rates were 45% and 65%, respectively. The

April-July period, during which scent-staticn visitation was

lowest, corresponded to the season of denning and rearing of

young, a period when tue adult fox population was at its

annual low and total daily acvements (2235 m) were minimal

(Progulske 1982). The subsequent peak in scent-station

visitation during the late-summer and fall season resulted

from an increase in population density due to the addition

of young, dispersal of juvenile foxes, and accelerated daily

movements of adults (4385 m) (Progulske 1982). However,

there were no significant differences among visitation rates

for these periods (P > 0.05).











Gray fox scent-station visitation rates were affected by

both population density and movements; however, population

density was probably the more important factor. To

illustrate, during the December 1S80-March 1981 breeding

period, total daily movements (8242 m) were nearly double

that of any other period (Prcgulske 1982); however,

scent-station visitation was intermediate during this same

period. Population abundance also was very likely

intermediate during the December-March period because both

juvenile dispersal had ended and over-winter mortality had

occurred. Therefore, scent-station indices for gray foxes

accurately reflected seasonal trends in population density,

despite seasonal changes in movement patterns.

Opossum population density, based on 166 captures of 60

individuals, was 10.1 per km2 during December 1980-March

1981. The scent-staticn visitation rate during this period

was 7.5%. The seemingly erratic trends in scent-station

visitation rates by opossums (Fig. 1), as well as the

significant interaction between year and month (Table 1),

indicated that scent-station indices probably did not

reflect trends in opossum population abundance.















CONCLUSICES ANE RECCHEE1DATICNS


Scent-station transects have been used widely for

indexing the population trends of carnivorous mammals

without an independent verification of population abundance

(Wood 1S5S, Linhart and Knowlton 1975, Brady 1981, Sumner

and Hill 1980, Morrison et al. 1981, Humphrey and 2inn 1982,

and others). In this investigation, scent-station transects

provided a reliable index of the population abundance of

bobcats, raccoons, and gray foxes, tut prctably not of

opossums. Yet, despite the utility of the technique,

differences in operational methodology, such as the spacing

of transects and stations within transects, frequency of

operation, tracking surfaces, and type and presentation of

attractant, have minimized the comparability of

species-specific visitation rates among different geographic

regions. Therefore, standardization and verification of the

technique must be achieved to facilitate inter-area

comparisons.

Findings from this investigation, coupled with previously

published works, facilitated the formulation of some

guidelines for standardizing the scent-station technique,

for obtaining annual trends in the abundance of mammalian











carnivores. First, transects should be operated when

visitation is highest, as recommended by Roughton and Sweeny

(1982). Second, transects with stations spaced at 0.32 km

were demonstrated to be indicative cf trends in the

population abundance of bobcats, raccoons, and gray foxes;

therefore, this interval of station placement appears

feasible when indices for multiple species are to be derived

from the same transect network. Third, transects should be

distributed so as to proportionately sample all major

habitat types, particularly when indices for the raccoon are

required; replicate transects per habitat-type are required

if habitat use is to be measured. The area-wide

distribution of transects, i.e., the number of transects per

unit area, required to accurately reflect population trends

remains unknown. However, the characteristic home range

size and movement pattern of individual species are probably

the most important factors in determining appropriate

transect placement and spacing. Fourth, transects operated

for 1 night only will provide reliable indices for abundant

mammalian species (Roughtcn and Sweeny 1982, unreported data

from this project); however, multiple-night sampling should

be considered for rare or uncommon species. finally,

although not specifically addressed in this project,

tracking surfaces and attractants should also be

standardized. Stations should be constructed with the test

available natural material to minimize the possibility of











the tracking surface becoming an attractant or repellent.

The attractant and the tethcd of presentation should be

standardized as suggested by Roughtcn and Sweeny (1982).

This study provided 2 specific recommendations for

operating scent-station transects in Florida, if indices for

all carnivorous furtearers are to be developed from a

transect network that is operated only 1 night per year.

First, transects should be operated in October or November.

Second, a large proportion cf the transects should be

distributed in wetland habitats. If manpower and time

constraints permit transects to be operated more than once

per year, bobcats and gray foxes should be indexed in

November and raccoons in September. If transects are to be

targeted specifically for bobcats, the optimal station

spacing should be investigated further. The

characteristically large home ranges cf bobcats during

periods of low population abundance (Erogulske 1982)

indicates that more bobcat home ranges could be encountered

by increasing the transect length.















LIfERATURE CITED


Brady, J. E. 1981. Preliminary results of bobcat
scent-stations in Florida. Pages 101-103 is Bobcat
research conference proceedings: current research on
biology and management of Lynx rufus. Natl. ildl.
Fed. Sci. and Tech. Ser. 6.


Conner, M. C. 1982. Determination of bobcat (JLnx rufus) and
racccon (Procyon Iotor) population abundance by
radioisotope tagging. M.S. Thesis. Univ. Florida,
Gainesville. 55pp.

Davison. R. P. 1981. The effects of exploitation on some
parameters of coyote populations. Ph.D. Eissertation.
Utah State Univ.. Logan, 153pp.


Hon, T. 1981. Relative abundance of bobcats in Georgia:
survey techniques and preliminary results. Pages
104-106 in Bobcat research conference proceedings:
current research on biology and management of Lynx
rufus. Natl. Will. Fed. Sci. and Tech. Ser. 6.


Humphrey, S. E., and T. L. Zinn. 1982. Seasonal habitat use
by river otters and everglades mink in Florida. J.
Nildl. Manage. 46:375-381.


Johnson, N. F. 1981. Efforts to understand Kansas' bobcat
populations. Pages 37-39 in Ectcat research conference
proceedings: current research on biology and management
of Lynx rufus. Natl. Vildl. Fed. Sci. and Tech. Ser.
6.


Johnson, K. G., W. G. Minser, III, and M. E. feltoa. 1981. A
survey of procedures used for indexing population
trends of furbearers in the southeastern United States.
Dept. of For., Wildl., and Fish., Univ. Tennessee,
Kacxville. 5pp. mimeo.











Knowlton, F. F., and W. M. Tzilkowski. 1981. Trends in
bobcat visitations to scent-station survey lines in
western United States, 1972-1978. Pages 8-12 in Bobcat
research conference proceedings: current research on
biclcgy and management of Lynx rufus. Natl. Wildl.
Fed. Sci. and Tech. Set. 6.


Laessle, A. M. 1942. The plant communities of the Welaka
area. Univ. Florida Biol. Sci. Ser. 4:1-143.


Linhart, S. B., and F. F. Knowlton. 1S75. Determining
relative abundance of coyotes by scent station lines.
Wildl. Soc. Bull. 3:119-124.


Morrison, D. W., R. M. Edmunds, G. linsccmbe, and J. W.
Goertz. 1981. Evaluation of specific scent station
variables in northcentral Louisiana. Proc. Southeast.
Assoc. Fish and Wildl. Agencies. In press.


National Cceanic and Atmospheric Administration. 1974.
Climates of the states. Vol. 1. Water Inf. Cent., Inc.,
Port Washington, N.Y. 486pp.


Pimlott, D. H., J. A. Shannon, and G. B. Kolenosky. 1969.
The ecology of the timber wolf in Algonquin Provincial
Park. Ontario Dept. Lands and For. 92pp.


Progulske, D. R., Jr. 1982. Spatial distributions of bobcats
and gray foxes in eastern Florida. M.S. Thesis. Univ.
Florida, Gainesville. 63pp.


Richards, S. H., and 8. L. Hine. 1953. Wisconsin fox
populations. Wisconsin Conserv. Dept., Tech. Wildly.
Bull. 6. 78pp.


Roughton, R. D., and M. W. Sweeny. 1982. Refinements in
scent-station methodology for assessing trends in
carnivore populations. J. Will. Manage. 46:217-229.


Schultz, D. 1979. Timber and soil type map of the Research
and Education Center, Welaka, Florida. Inst. of Food
and Agric. Sci., Univ. Florida, Gainesville. 122pp
minec.







80



Sumner, P. W., and E. Hill. 1980. Scent-stations as
indices of abundance in some furtearers of Alabama.
Prcc. Southeast. Assoc. Fish. and Wildl. Agencies.
34:572-583.


Veno, P. A. 1976. Successional relationships of five Florida
plant communities. Ecol. 57:498-508.


Wood, J. E. 1959. Relative estimates cf fox population
levels. J. Wildl. Manage. 23:53-63.






































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