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Population estimates, habitat requirements, and landscape design and management for urban populations of the endemic Big Cypress fox squirrel (Sciurus niger avicennia)

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Title:
Population estimates, habitat requirements, and landscape design and management for urban populations of the endemic Big Cypress fox squirrel (Sciurus niger avicennia)
Creator:
Ditgen, Rebecca Selfridge
Publication Date:
Language:
English
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ix, 132 leaves : ; 29 cm.

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Subjects / Keywords:
Animal nesting ( jstor )
Female animals ( jstor )
Golf courses ( jstor )
Landscapes ( jstor )
Pine trees ( jstor )
Squirrels ( jstor )
Street trees ( jstor )
Trees ( jstor )
Understory ( jstor )
Vegetation ( jstor )
Dissertations, Academic -- Wildlife Ecology and Conservation -- UF ( lcsh )
Fox squirrel -- Habitat -- Florida ( lcsh )
Wildlife Ecology and Conservation thesis, Ph. D ( lcsh )
Genre:
bibliography ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis (Ph. D.)--University of Florida, 1999.
Bibliography:
Includes bibliographical references (leaves 125-131).
General Note:
Printout.
General Note:
Vita.
Statement of Responsibility:
by Rebecca Selfridge Ditgen.

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POPULATION ESTIMATES, HABITAT REQUIREMENTS, AND LANDSCAPE
DESIGN AND MANAGEMENT FOR URBAN POPULATIONS OF THE ENDEMIC
BIG CYPRESS FOX SQUIRREL (Sciurus niger avicennia)













By

REBECCA SELFRIDGE DITGEN












A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA

1999



























Copyright 1999

by

Rebecca Selfridge Ditgen



















Dedicated to my father,
who first took me outdoors and shared his love for wild things,
and in loving memory of my mother,
whose life of gentle strength and patience continues to inspire me.















ACKNOWLEDGMENTS

I thank the Florida Fish and Wildlife Conservation Commission, Nongame Wildlife

Division, for their continuing attention to the pressures and consequences of rapid

landscape change in southwest Florida. I am grateful for their funding and the support

which made this project possible. I thank the staff of the Florida Museum of Natural

History for administration of the funding.

I thank my committee for their time and valuable insights. Dr. Stephen Humphrey has

given greatly appreciated support and encouragement, and with patience and generosity

has been the advisor I wanted and needed. Dr. John Eisenberg has shared his rich

knowledge of mammals and, as teacher and friend, has inspired me to reach for the best in

myself Dr. Mel Sunquist has enthusiastically shared his skills in fieldwork, his hallmark

sense of humor, and the joys of field classes at the Ordway. Dr. George Tanner has

provided an excellent balance between attention to detail and the long view, as well as the

important sense of perspective we often forget in the midst of all of this. And Dr. Mark

Brown has been gracious in his willingness to put up with an idealistic wildlife biologist.

My work relied on access to the private property of over 60 golf courses in Lee and

Collier counties. I thank the golf course superintendents, staffs and members of these

courses for their generous contributions to my work. Special thanks and deep appreciation

goes to the 2 clubs that facilitated the radio-telemetry studies centered on their property.

At Royal Poinciana Golf Club, Gary Grigg, CGCS, and his staff provided assistance and


iv








access for 2 years of work, while Dale Walters, CGCS, and his staff at Royal Palm County

Club assisted in my work for the final year of the field study. Courses adjoining these main

sites also permitted frequent visits to locate collared fox squirrels. These clubs included

Hole in the Wall, Wilderness, Country Club of Naples, Quail Run, Bear's Paw, Hibiscus,

and Royal Wood. I would also like to thank Tim Hiers and Mike Mongoven for their

contributions to my understanding of the management and history of golf courses in

southwest Florida.

Hardy thanks go to colleagues in the field and office. Joys and challenges of

trapping and collaring were shared by 2 excellent field assistants, Audrey Grieser and John

Pamilio. Jay Harrison helped with the analysis of the data on the 60 course sites. John

Shepherd gave generous assistance with vegetation sampling and mapping, and with GIS

quagmires. For lively discussions, timely and invigorating e-mail, and a shared passion for

work in the mammal world, thanks go to Lisa Molloy, Susan Walker, Deb Jansen, Kae

Kawanishi, and Candace McCaffery.

I thank the 38 fox squirrels whose fondness for peanut butter and pecans led them

to carry radio-transmitters for up to 20 months as they moved, mated, fought and played.

For support, understanding, and the courage to marry a woman in the final months

of writing her dissertation, I thank my loving and wonderful husband, John.













v















TABLE OF CONTENTS



ACKNOWLEDGMENTS ................................................. .................iv

A B STR A C T ........................................................... .. ........ .......................... viii

INTRODUCTION............................................................................................. 1
Fram ew ork .................................................................... .............................. 1
Problem ................................................................. ..... .......... ..................... 3

ST U D Y A R E A ...................................................................... ............... ............... 7

L andscape Study ............................................................................. ....................... 9
Radio-Telemetry Study ................... ............. ........ ................................................... 9

METHODS . .................................................... .............................................................. . 17

Landscape Evaluation and Censusing ............................................................ 17
Sam pling ......................................................... ................... .. . 17
Data Analysis ........................................................................................ 19
Habitat Use and Demography ......................................................... ...................20
Radio-telem etry ............................................................................. ................... 20
Vegetation Sampling .......................................................................................... 24
D ata A nalysis.................................................................................. .................... 26

R E SU L T S ........................................................................... .......................... ... ..... .30

Landscape Evaluation and Censusing ........................................................................... 30
Squirrel C ounts....... ............... ......................................................................... 30
Course A ttributes..................................................................................................... 30
Landscape Evaluation Index............................................................................ 34
Demographics and Habitat Use ..................................... 37
Survival ............................................................................................................. 37
R eproduction ............................................................... ......... ....................... 4 1
M ortality and D isease ............................................................................................. 42
P opulation E stim ates.................................................... ................... ..................... 43
H om e R ange................................................................................................... 43


vi









N est Sites .......................................................... ............. . .............. ............... 72
Landscape Composition and Vegetation Mapping ..................................................73
H abitat U se............................................................................................. 82
Feeding Patterns ................................................ ....................... .......................... 88

D ISC U SSIO N ..................................................................... ........................ ..... 94

Higher Quality Golf Course Landscape ...................................... ....................... 94
Lower Quality Golf Course Landscape ...................................... 101
Landscape Evaluation ....................................................................... ........................ 104
Sum m ary ..................................................................... ................... ....................... 110

MANAGEMENT RECOMMENDATIONS ................................. ................... 112

APPENDIX A, TREE SPECIES ON STUDY SITES ................................................. 117

APPENDIX B, GOLF COURSES, SQUIRREL LEVELS, AND LEI RANKINGS.... 119

APPENDIX C, HOME RANGE DATA FOR SITES 1 AND 2 .................................. 121

LIST OF REFEREN CES....................................... ........ ............................ ............... 125

BIOGRAPHICAL SKETCH ...................................................................................... 132




























vii















Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy

POPULATION ESTIMATES, HABITAT REQUIREMENTS, AND LANDSCAPE
DESIGN AND MANAGEMENT FOR URBAN POPULATIONS OF THE ENDEMIC
BIG CYPRESS FOX SQUIRREL (Sciurus niger avicennia)


By

Rebecca Selfridge Ditgen

December 1999


Chairman: Dr. Stephen R. Humphrey
Major Department: Wildlife Ecology and Conservation

The Big Cypress fox squirrel (Sciurus niger avicennia) is endemic to open forests

of southwest Florida. Rare in wild lands of southwest Florida, its remains on certain golf

courses in Lee and Collier counties. This study was carried out from August 1995 to

December 1997 to document squirrel population levels in a variety of golf course

landscapes, to determine habitat use and requirements of course populations, and to

provide guidelines for favorable landscape design and management in developing areas.

Elements of vegetation composition and structure, and landscape configuration

were recorded at 60 golf courses. Counts of squirrels were made at each course to

determine fox squirrel population levels. A Landscape Evaluation Index, developed from





viii








cluster and factor analyses of landscape elements, allowed ranking of the 60 courses in

terms of their suitability for fox squirrels.

Radio-telemetry was used to examine home range size, habitat use, and population

dynamics at 1 high quality course and 1 lower quality course. Tracking studies indicated a

density of 42.4-49.8 squirrels/km2 at Site 1 and a density of 6.3-8.2 squirrels/km2 at Site 2.

Squirrels fed heavily on pine and cypress from late summer to mid-winter, and relied on

native and exotic species between March and May. At Site 2, squirrels showed a heavy

reliance on feeders between January and July.

The Index identified 7 courses with high quality landscapes. All were part of 36

hole courses, contained large stands of open pine and cypress, and had large contiguous

areas free of automobile traffic. The remaining courses had unfavorable landscape

elements: isolation within developed landscapes, small stands of undesirable species, heavy

understory vegetation, and complex development patterns. Twenty-three courses offer

little opportunity for habitat improvement. Thirty courses can improve habitat for present

fox squirrel residents, but do not contain the landscape features required for long-term

populations.

Landscape design and placement are crucial in creating and preserving fox

squirrel habitat. Courses, or groups of courses, must contain large areas free of roadways

and development to allow safer movement within large home ranges. Vegetation must

include large stands of pine, cypress, cabbage palms, and associated native trees with

open understories.






ix
















INTRODUCTION


Framework


In the past decade, questions of ecologically sound landscape design and

management have attracted the attention of an increasing diversity of scholars and

practitioners. Researchers in wildlife and conservation biology, landscape architects,

planners, developers, and a range of physical and biological scientists have worked to

integrate a growing body of biological principles and physical science with species and

community ecology to manage landscapes for diversity and sustainability (Forman 1995,

Saunders et al. 1991, Soule and Kohn 1989, Turner 1989). With an understanding of the

rapid rates of land conversion and a cognizance of perceived human needs, the visions of

design and management reach from large-scale regional preserves (Carr et al. 1994) and

statewide, single species management plans (Mech 1998, Stith et al. 1996), to smaller

scale, county-wide urban parks (Mazzotti and Morgenstern 1997) and woodlot planning

(Fitzgibbon 1993).

Within this larger movement toward ecological design and management, stirrings

of interest have emerged in some previously untouched arenas. One of these is in the

community of golf course designers and managers, where individuals have begun to search

for ways to create more ecologically responsible golfing developments (Grigg 1990, Foy

1989). Golf courses, considered by opponents of development to be the antithesis of


1






2


ecological diversity and sustainability, have been portrayed as destroyers ofwildlands and

conduits for pesticides and fertilizers (Edmundson 1987, Foy 1989, Tietge 1992). In

response to such criticism and because of personal and professional interest in a more

diverse environment, advocates for change are attempting to move golf courses and

accompanying developments toward more ecologically sound designs, to create more

"naturalistic landscapes", and thus to encourage the preservation of native plants and

animals (Dodson 1990, 1994, Leuzinger 1994).

To date, much of the effort toward responsible golf course design and management

has focused on badly needed reviews of turf systems and course facilities, as well as

broader looks at maintenance of native vegetation in roughs (Balough and Walker 1992,

European Golf Association Ecology Unit 1995, Weston 1990, 1994). Scientific research

examining the benefits to wildlife of more natural and diverse golf course landscapes is just

emerging. Initial research addresses the role these newer or more "naturalistic" courses

may play as habitat for birds. Early work indicates they may contain more species than

surrounding landscapes of agriculture or dense development (Terman 1997). Because

even the most sensitively designed golf courses will save native vegetation in patches

separated by manicured, exotic grass, and generally contain private homes and

accompanying vehicle traffic, they cannot be expected to favor bird species easily

disturbed by humans or those in need of large areas without edges (Moul and Elliott 1994,

Terman 1997). Recent work in Kansas has shown that more natural courses preserving

large remnants of native vegetation may create an avian habitat intermediate to dense

development and wildlands (Terman 1997).






3


While there is a growing interest in understanding and improving golf courses as

habitat for avian species, little research has been undertaken to examine their ability to

support mammals or other non-avian wildlife, nor has research considered the impact of

surrounding landscapes on golf course wildlife (Terman 1997). Work by Jodice and

Humphrey (1992) in southwest Florida suggested that the threatened Big Cypress fox

squirrels may occur at higher densities on golf courses near the coast than in preserve

lands to the east (Jodice and Humphrey 1992, 1993, Maehr 1993). An investigation of

golf courses as potential habitat for these relatively small, though wide-ranging mammals,

offers an excellent opportunity to address the broader question of golf course landscapes

as habitat for non-avian wildlife.

Problem


Fox squirrels (Sciurus niger) are a diurnal, arboreal species inhabiting open forests

of the eastern and central United States (Hall 1981). The 4 subspecies of the southeastern

states are larger and more varied in color than those to the north and west and prefer open

pine forests with oaks and associated hardwoods (Kantola and Humphrey 1990, Moore

1957, Weigl et al. 1989). Of these, the Big Cypress fox squirrel (S. n. avicennia) is the

most restricted in geographic range, found only in the southwest tip of Florida, south of

the Caloosahatchee River and west of the true Everglades. Native to open stands of slash

pine (Pinus elliottii), cypress (Taxodium spp.), and tropical hardwoods, these squirrels

frequently feed and move on the ground. Their relatively large size and habits of ground

use make them especially vulnerable to the widespread landscape changes promoted in

recent decades (Humphrey and Jodice 1992, Moore 1956, Williams and Humphrey 1979).






4


Human activities affecting fox squirrel populations are widespread and varied in

southwest Florida. Changes in fires cycles on large preserves and privately owned forests

have allowed development of heavy understory vegetation not conducive to fox squirrel

movement and ground feeding. Conversion of range lands to citrus groves in northern and

central agricultural areas has eliminated open, parklike habitat favorable to fox squirrels

(Pearlstine et al. 1997). Rapid urbanization of coastal property in Lee and Collier counties,

from Naples to Ft. Myers, has created fragmented habitat with serious obstructions to

squirrel movement, resulting in isolated populations amid shrinking green space (Moore

1954, Williams and Humphrey 1979, Jodice and Humphrey 1993).

Demographic trends in Lee and Collier counties, two of the fastest growing

counties in Florida, clearly illustrate the forces driving land conversion in the coastal zones

of both counties. The 1970 permanent population of Collier County was 38,040. Between

1980 and 1990, Collier County grew by 77% (Collier County DESD 1996). In 1995,

Collier had a permanent population of 197,400 and a seasonal population of 245,000.

Projections indicate it will grow to between 508,00 and 770,00 by 2020 (Collier County

1996). The 1999 population of Lee County is given as 410,000 and is expected to reach

940,800 in 2020 (Lee County DCD 1998). Most of the development in both counties will

be concentrated along the coast, with Collier County expecting full development west of

highway 951 by 2050 (D. Weeks, person. commun.).

While fox squirrel populations have apparently declined in preserves such as Big

Cypress and Corkscrew Swamp and have vanished from dense housing developments and

commercial areas, they remain on certain golf courses within and near the burgeoning

developments of western Lee and Collier counties (Deborah Jansen pers. commun., Jodice






5


1990, Jodice and Humphrey 1992, 1993). Golf courses with remnant open pine and

cypress stands preserve fragments of suitable habitat within a swirl of traffic and

commerce.

In an effort to understand the ecology of these golf course fox squirrel

populations, Jodice (Jodice and Humphrey 1992) undertook a study of their diet and

activity patterns on 4 Naples courses in 1989-1990. His work relied on focal-animal

sampling of individuals located visually rather than by radio-telemetry, undoubtedly

allowing bias in a species that is often difficult to see. His work successfully highlighted

questions of population levels and suitability of golf course habitats (Jodice and Humphrey

1993, Maehr 1993). It became clear that little is known about spatial needs, movements,

habitat requirements, or demography of these urban populations and still less about the

prevalence of these golf course populations and the landscapes that might promote their

survival.

To evaluate the usefulness of golf courses as refugia for Big Cypress fox squirrels

we must know more about the ecology of the species as well as the impact of landscape

features and configuration on their survival. Can they feed, move, reproduce, and survive

for sustained periods within the fragmented habitats found on golf courses? Are all

courses suitable or just a few? And why? This study will address these questions by

looking at home range size, habitat use, feeding patterns and demography of fox squirrels

on golf courses and by identifying golf course landscape features favorable to their

survival.

The specific goals are as follows:






6


1. To survey the status of golf course populations and the landscape elements of a

range of golf course types in order to determine which features favor their survival. Sixty

courses in western Lee and Collier counties will be considered.

2. To gather data on home range size, dispersal, habitat use, and population

dynamics through the use of radio-telemetry. Two golf course populations in Collier

County will be studied, one with high numbers of squirrels and one with lower numbers of

squirrels.

3. To evaluate the potential of the 60 golf courses as refugia for urban populations

of fox squirrels.

4. To provide recommendations for design and management of golf course

landscapes to improve habitat for Big Cypress fox squirrels.














STUDY SITE

All study sites were located in the western half of Collier and Lee counties in

southwest Florida (Fig. 1). The area has a humid subtropical climate with heavy influence

from the surrounding warm waters and the seasonal changes in the Bermuda high (Chen

and Gerber 1990). These features give rise to cool dry winters and warm, rainy summers

and autumns, with extreme events such as occasional hard frosts and hurricanes playing a

strong role in the composition of the vegetation community. Native vegetation of the

flatwoods physiographic region in which the sites were located includes pine flatwoods,

cypress domes, and mangroves. The presence ofEntisols, Histosols, and Spodosols

reflect a mixed terrain of high, relatively dry, sandy ridges, and low, poorly drained

swamps (Brown et al. 1990).

Patterns of temperature and precipitation varied from year to year in the 3 calendar

years of the study, with wide deviations from normal in summer precipitation (NOAA,

1995-97). Summer and fall of 1995 were extremely wet. Stations at Ft. Myers and Naples

reported 1.7 m or more of precipitation between June 1 and October 31, more than 0.76 m

above normal. Flooding and long-term standing water were common on most sites during

late summer and fall 1995. In the same months of 1996 the stations received only 0.56 m

of rain, a 0.36 m deficit. January 1997 to August 1997, when the tracking studies ended,

had normal levels of precipitation. The winter of 1995-1996 had at least 2 cool periods,

with 4 nights of0 C , and widespread damage to the more tropical flora. The winter of


7





8









\ / *1 \Collier












Site
SSouth
- � 'Florida







! "




S-75

o Site 2







Marco
Island






Figure 1. Location of study sites in western Lee and Collier counties. Black
dots are clubs visited for landscape and squirrel surveys. Red dots are sites
of the radio-telemetry studies.






9

1996-1997 was warmer than normal. An average January, with warm weather (19 C) and

1 light frost, was followed by 2 months of high temperatures. February averaged 22 C, 3.5

C above normal, and March 24 C, 3 C above normal.

Landscape Study


Of the 60 golf courses selected for the landscape analysis and fox squirrel

censusing, 18 were in Lee County, south of the Caloosahatchee River, and 42 were in

Collier County. Course landscapes ranged from undeveloped, with large tracts of native

vegetation, to intensely developed courses having close-set, multiple story condominiums

on both sides of the fairways. Highly developed courses usually allowed for few trees,

native or exotic, in the roughs. Courses ranged in age from over 40 years to those recently

opened and still under development. The oldest courses in the study, located near the Gulf

Coast, commonly were isolated from other clubs and were surrounded by development.

On the eastern edge of development courses tended to be newer, often grouped together,

and were located within a mixture of increasing development and remnants of pine and

cypress stands.

Radio-Telemetry Study


Two 18-hole golf courses in Collier County, Florida, were selected for the radio-

telemetry studies (Figs. 2,3). Site 1, the 18-hole Royal Poinciana Cypress Course, was half

of a 36-hole private Royal Poinciana Golf Club built in 1971 in central Naples. Royal

Poinciana has no residential development within the 135 ha of the golf course grounds.

Fairways are bordered by open stands of moderate-size pines (Pinus elliottii var. densa),





10



Pine Ridge Blvd



QUAIL RUN

COUNTRY
CLUB OF
NAPLES









S HOLE IN CL
THE WALL






WILDERNESS




N

Golden Gate Parkway
BEARS PAW


Figure 2. Site 1, location map. Identification of golf courses known to be used
by fox squirrels from Site 1, Royal Poinciana Golf Club, Cypress Course.
Five courses adjoin Site 1, including the Pines Course in the same club.
Bear's Paw is south across Golden Gate Parkway. Stippled area is pine forest.























































0 .5

Figure 3. Site 1, aerial photograph. Configuration of Site 1 and the
surrounding landscape. Royal Poinciana Cypress Course is located in the
center, with 6 other 18-hole courses nearby. Names on Fig. 2.






12


cypress (Taxodium ascendens and T. distichum) and cabbage palms (Sabal palmetto), and

plantings of non-native broad-leafed evergreens (Appendix A).

Automobile traffic around the Cypress Course is limited to a short segment of

private entrance roadway on the north side of the Cypress Course, with no public

roadways on the boundaries of the course. Three golf courses, including the 18-hole Pines

Course within the same club, comprise the western, northern and part of the eastern

boundaries. The south boundary is an undeveloped pine stand of 115 ha and the

remaining eastern boundary is residential development of varying density. Royal Poinciana

and the neighboring clubs are located within a tract of approximately 1020 ha containing

6, 18-hole golf courses, of which 3 are undeveloped, 2 moderately developed, and 1

heavily developed. The tract contains 230 ha of forested land, ranging from drier pine to

swampy cypress stands. The 1020 ha tract is bordered by 4 extremely busy roadways,

Goodlette-Frank, Pine Ridge, Airport-Pulling, and Golden Gate Parkway.

Known predators at the site included eagles, bobcats, great horned owls, raccoons,

rat snakes, and the club house cat, which was allowed to roam the course at night.

Site 2, Royal Palm Country Club, is a developed 18-hole course near the eastern

limit of intense suburban development along Highway 41 East (Figs. 4 & 5). The club,

built in 1970, and the adjoining housing development cover 150 ha, of which 75 ha are

private homes, condominium property and roadways. The club lies within a landscape

currently undergoing rapid and dense development. All the fairways have development on

both sides, either single family homes or large condominiums. When I began research at

this site in 1996, 8 undeveloped, pine-covered lots remained along the fairways. In the

next 12 months at least 4 of these were developed, with all of the pines being cut. The











ROYAL
WOOD


Rattlesnake Hammock Rd.







HIBISCUS






St. A ndrews Bld .


Figure 4. Site 2, location map. Royal Palm Country Club, Site 2, and 2 neighboring 18-hole courses.






































KM

0 .5
Figure 5. Site 2, aerial photograph. Royal Palm Country Club, and 2 neighboring 18-hole courses. Names on Figure 4.






15


dominant vegetation of the Royal Palm roughs and the surrounding condominiums is

open pine stands. Non-native trees of a limited variety are scattered around the course and

are common in the small lots of the surrounding private homes. Cypress trees are present

but are not common and generally are small. The southwest boundary of Royal Palm

adjoins the public 18-hole Hibiscus Golf Course. The Hibiscus course and surrounding

dense development, together 132 ha, have few native tree species and narrow, open

roughs with only small and scattered stands of trees (Appendix A). Busy 2-lane and 4-

lane roadways border Royal Palm on the north, east and south, and Augusta Boulevard

bisects the course from north to south. Known predators at the site included eagles,

raccoons, and a domestic cat.

Activity on the courses changed with the seasons. Golf play was heavy on both

courses from December until April, with Royal Palm frequently having play on every hole

from 0730 until late afternoon. Royal Poinciana was generally less crowded. Summer play

was light, with each course closing 1 day a week for intensive maintenance work on the

course and roughs. Daily maintenance work began at 0530 or 0600 and continued until

late afternoon. In winter, maintenance crews worked on the courses every day, 0630 until

1430 weekdays, and 0630 until noon on weekends. Maintenance for the removal of

vegetation in the understory of tree stands included mowing, hand removal of shrubs and

herbaceous plants, the addition of pine straw and the use of herbicides. Both clubs

irrigated the fairways, greens and tees in the early morning and often again in the early

evening.

Cone production on the study courses was noticeably higher in 1996 than 1997. In

the summer and fall of 1996, following a wet 1995 summer and fall, cone production on





16


both pine and cypress trees appeared to be high, with both species heavily laden

throughout the Cypress Course. In 1997, little cone production was evident on pines and

the cypress suffered an infestation of tent caterpillars in the late spring, resulting in

widespread defoliation of cypress and lower early season cone production.














METHODS


Landscape Evaluation and Censusing


Surveys of golf course landscapes and fox squirrel populations were conducted to

determine the status of fox squirrels and to identify landscape features favorable to their

survival.

Sampling

Sixty golf courses in western Lee and Collier counties were selected for landscape

analysis and squirrel censusing. In selecting the courses I looked for a range of landscape

types relating to

* density of development surrounding the club boundaries, from heavily developed to
undeveloped;
* type and configuration of development on the course, from undeveloped,
nonresidential courses to those with dense development of houses and/or
condominiums;
* character of the rough vegetation, from little and scattered, through open tree stands,
to dense forest stands with heavy understory.

To obtain pre-visit information about age and landscape configuration for many of

the 80+ courses in Lee and Collier counties I interviewed Tim Hires, Collier's Reserve

Country Club, Naples, and Mike Mongoven, Ft. Myers and Eastwood Golf Clubs. I

selected 66 courses for examination and contacted the courses through their

superintendents of maintenance, individuals frequently most familiar with the landscape





17






18


activities and wildlife on the courses. Superintendents proved to be excellent, and often

interested, sources of information and influence.

I made introductory visits to a majority of the 66 courses in August 1995-

December 1995. Exceptions were made for courses flooded by heavy summer and autumn

rains. Three clubs did not wish to take part in the study; from the remaining 63 courses I

selected 60 courses in 47 clubs, 42 in Collier County and 18 in Lee County (Appendix B).

Each club in the study was visited 3 times for censusing fox squirrels and

landscape analysis. The exceptions were those courses at which I did not see fox squirrels

or sign of fox squirrels on the first visit and the superintendent and course workers had

not seen fox squirrels for at least 1 year. At these courses I made no more than 2 visits for

landscape analysis and squirrels censusing. Squirrel censusing took place between

September 15 and May 15 each year, as squirrels were less active on the ground in the

summer. Squirrel counts were conducted mornings, 0700-1030, or late afternoons, 1500-

1730, when squirrels were most likely to be on the ground and visible (Jodice and

Humphrey 1993). Sampling times were limited to sunny days with light or no wind and

temperatures over 180 C.

I sampled for squirrels by driving around the course in reverse order, to decrease

my interference with golfers. I stopped at each fairway for 10 minutes and selected 1 or 2

locations most likely to attract squirrels: open tree stands, trees with food items, and

feeder areas. I searched trees and areas with open ground while listening for sounds of

movement and communication. I recorded squirrel sign, nests, cone middens, and palm

leaf or bark pealing. I recorded each squirrel spotting and did not attempt to determine if

animals were recounted after moving to another area of the course. I counted individuals






19


on private property adjoining the fairways, such as those using feeders. The highest count

of the repeated visits was recorded as the number of squirrels seen at that course. Five

categories of squirrel counts were created to allow grouping of courses by population

levels. They were

* Level 1--none present, a course at which I did not see fox squirrels and the
superintendent and workers had not seen them for over a year;
* Level 2--none seen, a course at which I did not see fox squirrels but the superintendent
reported sightings within the past year. This often meant traveling squirrels were on
the course for a few weeks or 1 or 2 come from neighboring courses for occasional
use.
* Level 3--low, the highest number of squirrels seen on the course was 1-5.
* Level 4--medium, the highest number of squirrels seen on the course was 6-10.
* Level 5-high, the highest number of squirrels seen on the course was 11 or more.


In the evaluation of habitat variation between courses I was interested in

landscape features that could impact squirrel feeding, movement, nesting or predation.

With these needs in mind I collected data on course configuration, place in a larger

landscape, vegetation composition and structure, predators, course history and human

interactions with squirrels. A field survey of questions that could be answered with a yes

or no response was developed to report on this range of landscape attributes. All

questions could be completed during a 3 hour tour of an 18 hole course and a 20 minute

interview with a knowledgeable superintendent. All final landscape surveys were

conducted between April and December 1997.

Data Analysis

Fifty-one responses, or attributes, from the landscape surveys were used to

examine landscape variation among the 60 sampled courses. Two methods of examining

variation were used. Single linkage cluster analysis was used to identify aggregations of






20


similar courses based on 51 landscape attributes (Statistical Analysis System 6.0). Two

prominent clusters were selected for further examination. Chi-square (a=0.05) was used

to test the hypothesis that attributes were randomly distributed between these clusters.

Factor analysis was used as another method to identify factors or groups of attributes that

explained variation among the courses (SPSS).

The 29 distinguishing attributes identified though cluster and factor analyses were

used to create a Landscape Evaluation Index (LEI) (Bender et al. 1996, Brooks 1997,

Reading et al. 1996, Thomasma et al. 1991, USFWS 1980, 1981). The Index allowed

ranking of all courses and a comparison to squirrel population levels. These attributes

were grouped into 3 components according to the landscape feature they described: 1)

vegetation, 2) ground cover, and 3) landscape position. Attributes were weighted

according to the differences in their frequency between high and low cluster courses and

their ranking in factor analysis. For each course, the sum of attribute weights was

expressed as a fraction of the sum of weights for an ideal course. A score of 1 indicated

that a course had all of the desirable characteristics; a score of 0 indicated that it had none.

The geometric mean of the 3 component scores was taken as an overall LEI (Reading et

al. 1996). Courses were then ranked according to the Index and compared to patterns of

squirrel sightings, as indicated by the 5 levels of population (Van Home 1983).

Habitat Use and Demography


Radio-telemetry

A radio-telemetry study was conducted to gather data on home range, dispersal

patterns, habitat use, feeding patterns and demography of two fox squirrel populations.






21


Two sites were selected for the study, one without residential development, containing a

high number of squirrels, and another, well developed, with lower numbers of squirrels.

Criteria for selection required that the sites have: 18-hole courses with similar size

fairways and roughs, prominence of open pine stands, a course configuration that allowed

movement around the course throughout the day, absence of 3 highly invasive exotic tree

species (Casuarina spp., Schinus terebinthifolius, and Melaleuca quinquenervia),

squirrels that were not fed by golfers and therefore not tame, and the strong support of the

course superintendent and the club greens committee for the research project. The last

was especially important as the clubs would provide golf carts for 12-20 months of

tracking endeavors and would support the regular presence of a non-member researcher

and collared fox squirrels on their courses. Permission to work on Royal Poinciana

Cypress Course was granted in November 1995 and that for Royal Palm in July 1996.

Trapping took place during 4 periods: Site 1 only, December 1995-March 1996;

Sites 1 and 2, July-August 1996, November 1996-February 1997, and July 1997. Because

of the public nature of the trap sites and the desire to decrease stress to individuals,

especially females who might be pregnant or nursing, a trap line was not used. Instead,

squirrels were trapped using a focused trapping method in which 1 or more traps were set

for 1 or 2 specific individuals in a small area. One-ended Tomahawk #204 squirrel traps

were baited with an oily, natural peanut butter and pecans. Traps were placed on the

ground under trees where squirrels were feeding or resting or within 7 meters of

individuals feeding on the ground. Often the traps were covered with Spanish moss or

palm leaves. The traps were baited, set and covered with moss at some distance from the

trap site and rapidly dropped off from a golf cart. Squirrels were acclimated to carts and






22


would return to the place of feeding after I moved away from the trap. I watched the traps

from 30-50 m. In the season of low food supply, squirrels could frequently be baited into

the traps within 10 minutes to 1 hour. This method was generally successful during the

winter months, when 2 or 3 individuals might be trapped and collared in a day. In the

summer and autumn months, particularly in food-rich 1996, squirrels were extremely

difficult to trap.

To ensure recapture and collar removal at the end of the study in July 1997,

individuals were baited with oily peanut butter and pecans for 2 weeks prior to the

trapping period as they were located during normal radio-tracking. Final trapping was

further aided by an apparently lower pine and cypress productivity in 1997.

Trapped squirrels were covered and moved to the cart within 1 minute. Removal

of the trap cover encouraged them to move into a dark cloth and net restriction tube that

was attached to the opening end of the trap. While constrained in the bag they were

weighed and given an injection of Ketamine HCI (100mg/ml) in the hip. Individuals 675-

800 gms were given 0.25cc, those 800-1000 gms received 0.3cc. After 4 minutes or when

they showed little sign of movement, they were removed from the bag and were tagged in

both ears with monel sequentially number tags (size #3, National Band and Tag, Newport,

KY), measured, aged, fitted with radio-transmitters and photographed. Females with

darkened nipples of any size were considered adult. Males with developed testes

descended into the scrotum were considered adult. Males with no obvious scrotum

development or with slight development were considered subadult. There was a clear

difference in the pelage and scrotal development between subadult males who had never

developed sexually and adult males undergoing seasonal fluctuations of testicle






23


development. Subadults had shorter, fine fur and no vestige of scrotal development. No

animals under 5 months of age were captured.

Squirrels were released at the site of capture after spending 3-4 hours in a 60 x 30

x 30 cm ventilated wooden wake-up box. Rapid retrieval, covering of the trap and

immediate anesthetization appeared to reduce trauma; no squirrels died during trapping or

collaring procedures.

From December 1995 to mid 1996, 25 gm AVM (AVM Instrument Co.,

Livermore, CA) radio transmitters configured as resin pods with machine belting neck

bands and 6 inch back antennae were used. This model proved unsatisfactory due to

repeated transmitter failure, poor service and removal by squirrels cutting the belting. In

late 1996, I began using ATS (Advanced Telemetry Systems, Inc., Isanti, MN)

transmitters with resin pods, very fine stainless steel chain neck bands and back antennae,

total weight of 28 gms. These worked extremely well, with no radio failure or removal by

the squirrels. Final recapture did show that 2 individuals had slight neck abrasions.

Collared squirrels were located a minimum of 2 times a week, except when

weather, golf course conditions or course use would not allow (Mech 1983). Individuals

were located once a week in 2 of 3 daily tracking periods, 0630 to 1030, 1031 to 1430,

and 1431 to 1900 EST. Squirrels were frequently located more than once in a sampling

period; data were collected on each siting. Open vegetation and ready access to trees

allowed visual sighting following radio location. When a squirrel was in a nest or

concealed by heavy vegetation and visual sighting was not possible, I was able to identify

the tree and the area of the tree in which the animal was located. When an animal moved

to another golf course I used triangulation (White and Garrott 1990) to determine its






24


location and then I traveled to the course for visual sighting and collection of activity data

if possible. When a squirrel disappeared from the course and could not be located at a

neighboring course, I searched the surrounding area in all directions. In December 1996, I

conducted an aerial search of Collier County west of highway 951 in an attempt to locate

squirrels that had disappeared.

Once an individual was sighted, its location was mapped on aerial photographs.

Recorded data included: time, activity (3 points at 60 sec. intervals), nature of the site and

location at the site, food type if feeding on identifiable material, reproductive condition if

visible, number of squirrels present within 5 m (both fox and gray), and number of collared

squirrels within 5 m. Records of temperature, dew point, sky condition, and wind were

recorded at the start of each session.

At least once a month throughout the course of the study I took visual counts of

fox squirrels at each study site. I followed the procedure outlined for the 60 course

squirrel counts and in addition recorded if each sighted squirrel was collared or uncollared.

These counts were used to estimate the fox squirrel population of the 2 study sites.

Vegetation Sampling

The tree stands on three courses, Royal Poinciana Cypress and Pines and Royal

Palm, were sampled and mapped to allow comparison of habitats used by the two radio

collared populations. The Pines course was included in the vegetation sampling because it

bordered Site 1 on 2 sides and collared male squirrels frequently used the area. The large

forested stands at the Poinciana courses were sampled using a structured pattern of 20

meter diameter circular plots placed at intervals of 25 meter from center to center on a

north-south line and at intervals of 30 meters from center to center in an east-west line.






25


This arrangement was designed for the most complete yet rapid sampling of the generally

north-south tending forested areas. In the front 9 of the Pines course the pattern was

oriented in an east west direction as the fairways in that section ran at right angles to the

rest of the club. Within each plot, all trees over 10 cm dbh were identified and measured.

All palms were counted. Presence of all saplings were recorded. Understory coverage was

recorded as percentage and type and the ground cover was recorded as litter if over half of

the plot had a significant layer of pine or pine and cypress litter. If not noted, ground cover

was dominated by grass with occasional patches of bare soil.

In the narrow or small plots of Royal Poinciana and for the entire Royal Palm

course, all the trees in each discrete plot were counted. If 10 or fewer of 1 species were

counted, each tree was measured. If more than 10 of 1 species was found in a stand, 10 of

the trees were measured. Saplings and ground cover were recorded as in the circular plots.

The Royal Palm site contained a large region of private lands belonging to

condominium complexes and private homes. All pines and cabbage palms on the condo

lands were counted and all trees on private lands on the 3 streets of private housing were

counted.

Seven courses known to be used by the squirrels of the 2 study sites were

evaluated using the method designed for the 60 course landscape evaluation portion of the

study. This provided data on tree species present, identification of dominant species, types

of ground cover, proportion of the course in tree stands, density of the understory and

types of ground cover. The large pine stand south of the Cypress course and the

developing landscape east of the Pines course were not accessible for sampling.






26


Data Analysis

In January 1996, I conducted 10 repeat counts of fox squirrels on the Cypress

course to estimate the reliability of my squirrel census technique. The counts followed the

standard format presented in the landscape analysis methods section. Results of the 10

counts ranged from 8-16 with a mean of 12.4 and standard error of 0.79. Calculation of

95% confidence limits for a small sample gave a range of 10.6-14.2 squirrels (Fowler and

Cohen 1992). Such a range was considered reliable for the populations with higher

numbers of squirrels. Reliability in smaller populations was expected to be lower.

Survival and birth rates of collared fox squirrels were calculated on 6 month

intervals, as opposed to 12 months, to allow for movement of subadults out of the sample

population on the Cypress course or into the adult cohort of the sample population. Only

squirrels persistent in the sample population were considered survivors. Squirrels no

longer persistent in the sample population included subadults who dispersed to other

locations, individuals who disappeared, adults, generally males, who no longer used the

study site course but were known to remain on neighboring courses, and individuals

known to be dead. Survival rates for adult females, adult males, subadult males and

subadult females were calculated as the proportion of the collared squirrels remaining

active in the study site at the end of a 6-month period. The birth rate was taken as the

number of young known to leave the nest. The survival rate for juveniles was the

proportion of the summer 1996 cohort of young known to be alive at the end of 6 months,

7 of whom were part of the collared population in 1997. Most subadults, collared at

around 6 months of age, moved into the adult cohort at about 12 months of age.





27

Estimates of population size, the number of fox squirrels using the Cypress course,

were taken from 13 counts of collared and uncollared fox squirrels on the course taken in

the spring and summer of 1996 and spring of 1997. In late summer and fall 1996, counts

were not taken because the number of collared squirrels had declined due to collar

removal and squirrel sightings were extremely low during these months of concentrated

feeding in pines. Procedure for the counts followed that presented in the initial description

of fox squirrel censusing. From these counts I calculated the minimum know alive (MKA),

the Lincoln Index, and the Bailey unbiased population estimator for small sample sizes

(Bailey 1952, Krebs 1999.).

I used the RAMAS Ecolab software (version 2.0, Applied Biomathematics) to

estimate growth rates in the Cypress population. Estimates of the 6-month survival and

birth rates of life stages were used to construct the stage-based population projection

matrix. I used birth rates from the lowest and highest 6 month periods and an average of

the 3 periods to generate a range of growth rates seen during the study.

Tracking locations were digitized using Atlas GIS software (Strategic Mapping,

Inc., Santa Clara, CA) on an overlay of air photographs (TRW-REDI Property Data,

1996) registered to 7.5 minute topographic quadrangles. Accompanying data for each

point were coded and attached to point locations using Atlas GIS software.

Home range was determined using the kernel method and CALHOME software

(Kie et al. 1996, White and Garrott 1990, Worton 1989, 1995). Only 1 point per sampling

period was used in home range analysis (Cresswell and Smith 1992). Points were selected

randomly in cases where 2 or more points were recorded during 1 sampling period. The

95% contour was used to define the home range boundary of each individual and the 50%





28


contour as the core area (Kenward 1992, Wray et al. 1992). I calculated separate home

range data for year 1 (December 1995-October 31, 1996) and year 2 (November 1, 1996-

July 30, 1997) to allow for changes in the make up of the collared population. I used a

two-tailed t-test for paired samples (a=0.05) to compare home range sizes of Site 1

females that appeared in both year 1 and year 2. I compared home range size of adult

females to adult males in both years and 2nd year subadults from Sites 1 and 2 using a

Mann-Whitney test (a=0.05) (Fowler and Cohen 1990). The small number of adult

females and adult males at Site 2 precluded statistical comparisons using those individuals.

Habitat maps were created from aerial photographs using Atlas GIS software. All

vegetation sampling plots, including condominium areas, water features, residential areas

and streets, and course fairways and non-forest roughs were outlined and the area of each

type was measured. I calculated basal area, density, relative basal area, and relative density

of each tree species by plot. Nine categories were defined to represent the diversity of

vegetation seen on all 3 courses. Vegetation categories were defined by the relative basal

area (percent of total dm2/ha.) of pine, cypress, cabbage palm, and other native and exotic

species. Tree stands were then categorized by density (stems/ha.). Plots with a pine

needle litter layer or a shrub layer were identified and mapped as such.

Using Atlas GIS to analyze use of the vegetation types, tracking points were

overlaid on maps of vegetation classes for each course. I compared actual use of

vegetation types to that predicted by the percent area of each vegetation category. Chi-

square tested (a=0.05) the hypothesis that tracking points were randomly distributed

among vegetation categories.






29

Feeding patterns were examined by analyzing the tracking data in which the food

item was clearly identified. All food types taken more than 5 times in the 19 months of

feeding observations were considered. Changes in monthly feeding patterns were

determined by calculating the percent monthly total of each species or food type

consumed each month for the 19 months of feeding records. A diversity index of species

use, the inverse Simpson (Krebs 1999, MacArthur 1972, Williams 1964), was used to

measure both richness and evenness of use of the food types.














RESULTS


Landscape Evaluation and Censusing


Squirrel counts

I was able to sight fox squirrels at all courses that reported regular observations by

course personnel. During the squirrel counts, 5 or fewer squirrels were seen at 48 (80%)

of the 60 courses. Fourteen courses (23%) were level 1, with no squirrels seen during

surveys and no sightings by course staff in the past year. Nine courses (15%) were level 2,

with no squirrels sighted during the surveys, but course personnel reported occasional

sightings in the previous year. Reported sightings on these courses were frequently

traveling squirrels or an occasional visiting squirrel from a higher level neighboring course.

Level 3, 1-5 sightings, was the largest category with 25 courses, 42% of the total. I

sighted 6 or more squirrels on only 12 courses: 6 courses (10%) were level 4, 6-10

squirrels, and 6 were level 5, with more than 10 squirrels seen.

Course attributes

Cluster analysis of the 60 courses with 50 attributes produced a dendrogram with a

prominent cluster of 11 courses and a broader cluster of 18 courses (Fig. 6 ). The 11

course cluster, cluster 1, contained courses with a high occurrence of attributes favorable

to fox squirrels and the 18 course cluster, cluster 2, contained courses with a high number

of landscape attributes unfavorable to fox squirrels.



30













300- Cluster 2, 18 Courses

Cluster 1, 11 Courses

250




200



, I o
150-







0 -1 1 1 1 1 11 1 1 1 1 HI I I I I I 1 1JI
100




50






Figure 6. Dendrogram of single linkage cluster analysis. Arrangement of 60 golf courses using 50 attributes of vegetation,
ground cover, and landscape position.






32


Chi square tests of the 50 attributes showed that 20 attributes were non-randomly

distributed between the 2 clusters. Courses in cluster 1 were characterized by: large

patches of pine and cypress trees, open understories in the tree stands, large areas of pine

litter, having adjoining courses, being part of a multi-course club, and having at least 50

acres of adjoining forest. Cluster 2 courses were characterized by: a high degree of

isolation, few or no sizable tree stands, low numbers of pine and cypress, no obvious

dominant tree species, heavy development and busy roadways around the course (Table

1).

Factor analysis produced 2 independent factors or groups of attributes explaining

27% of the variation among the courses. Factor 1 accounted for 14.6% of the variance

and factor 2 for 12.3%. Factor 1 describes variation similar to that seen in cluster analysis,

large stands of open forest on larger, grouped courses vs well developed, isolated, courses

with more sparse tree cover and few native species. Thirteen attributes contributing to

factor 1 were the same as those showing non-random distribution between clusters in the

previous analysis (Table 1). Factor 2 is a comparison of "natural" courses with a heavy

understory to courses with clear understory and the presence of planted exotic trees. Eight

attributes contributed to the variation explained by factor 2 (Table 1).

While cluster and factor analysis were quite useful in identifying characteristics that

distinguished high and low quality courses in this study, it must be noted that they do not

provide a complete listing of all features that are either beneficial or harmful to squirrel

survival. Attributes such as untrimmed cabbage palms, while helpful to fox squirrels were

found on over half of the courses and occurred in all types of landscapes. Similarly, known

predators such as domestic cats and raptors were found throughout the






33


Table 1. Distinquishing attributes as selected by cluster and factor analyses. Column
1, factors which appeared non-randomly in high and low clusters. Number is the
percentage difference of occurrence between high and low clusters. Column 2, an
attribute in Factor 1 or Factor 2 as determined through factor analysis. Column 3,
attributes are grouped as components of the Landscape Evaluation Index. Column 4,
a desired answer indicates it is an attribute favorable to fox squirrels. Index weight
determined by columns 1 and 2.

1 2 3 4 5 6


SI2

M 5Attributes
Vegetation- well forested, large patches around course and
76.3 1 1 Y 7 11 often between fairways
72.7 1 1 Y 7 19 pines stands of 20 or more on most holes
81.8 1 1 Y 7 33 Pines or pines co-dominant
72.2 1 Y 5 17 over 100 large pines/18 holes
79.8 1 1 Y 5 18 pines occur throughout the course
61.1 1 1 Y 5 20 over 50 large cypress/18 holes
56.1 1 1 Y 5 21 at least 4 stands 20+ cypress trees/18hole
74.2 1 Y 5 29 Palmetto present
2 1 Y 3 30 4 or more exotic food trees common
course "tight", narrow roughs,trees scattered,
88.9 1 1 N 7 9 few or no large stands
85.4 1 1 N 7 34 no obvious dominant/s
2 1 N 5 13 3+ marginally managed stands/18 holes
heavy forest vegetation in roughs, can't see
2 1 N 3 12 more than 1 holehousing 0-iside
2 1 N 3 36 mixed natives majority of forest/stands
2 1 N 3 47 Large snakes present
50.0 1 N 1 31 eucalyptus present
Ground open understory, large areas pine litter, open
85.4 1 cover-2 Y 7 16 soil
2 2 Y 3 15 managed forests, understoy 75%+clear
unmanaged stands show vine invasion and/ or
2 2 N 3 14 other dense understory
Landscape
50.5 1 position-3 Y 5 5 club has 36 holes or more
1 3 Y 5 40 at least 1 adj. course has high levels F.S.
65.2 3 Y 3 39 adjoining forest over 50 acres
54.0 3 Y 1 51 10-25 years old in 1997
77.8 1 3 N 7 37 Completely surrounded by development
68.7 1 3 N 5 3 club has only 18 holes
50.0 1 3 N 5 41 0 adjoining courses with fox squirrels
70.7 3 N 5 43 1 or more boundaries with busy 2 or 4+ lane
2 3 N 3 50 less than 10 years old in 1997
38.9 3 N 1 38 0 adjoining courses





34

range of course landscapes. For this reason they did not appear as distinguishing attributes

of high or low clusters. The list of distinguishing characters must not be confused with a

more complete listing of characters favorable or unfavorable to fox squirrels.

Landscape Evaluation Index

The Landscape Evaluation Index was calculated using 29 attributes, 16 in the

vegetation component, 3 describing ground cover, and 10 relating to landscape position

(Table 1). The 60 courses ranged from 0.0 to 0.987 on the scale of 0.0 to 1.0 (Appendix

B, Fig. 7)). Seven courses were over 0.90, 18 from 0.50 to 0.751, and 35 were below

0.50. There was a gap with no courses from 0.90 to 0.752. Examination of the landscape

configuration of the courses shows that the 7 courses above 0.90 are either undeveloped

or have a perimeter development plan for each course or for the entire club (Fig.8).

Courses with a LEI of 0.75 or lower generally have more complex development plans

such as the shown in levels B and C in Figure 8. These require squirrels to cross streets or

travel through or around larger forest stands with dense understory vegetation to move

from one portion of the course to another.

Index values were strongly correlated with squirrel levels (r = 0.747, p<<0.01).

Courses that adjoin high level courses, but do not themselves have an index value over

0.90, are identified as Neighbors in Figure 7. Three of these courses have level 4

populations, though their index values would indicate a lower capacity, and 4 of the level

3 courses have higher populations than expected. Four of the level 1 courses have

moderate LEI ratings, yet no squirrels. These courses are newer, developing courses at

which the large forested areas have heavy, closed understory vegetation not favorable to

fox squirrels.














4 - 4 4 .0









�2 ** ** * *<



0 1 * i *







0
1.100 1.000 0.900 0.800 0.700 0.600 0.500 0.400 0.300 0.200 0.100 0.000
Landscape Evaluation Index
Figure 7. Comparison of Landscape Evaluation Index values to squirrel population levels. Trend line is a linear regression
(r2=0.557). Courses that adjoin the 7 courses (level 5 has 6 courses, symbols overlap) with index values of 0.90 or higher
are identified with gray diamonds as Neighbors. Four courses that have large areas of native tree stands with heavy
understory growth are identified by large gray squares, in level 1.






36







GolfCo use







Al. Perimeter development A2. Concentrated interior
development










SItm r B2. Expanded interior
development development with
preserved wetlands









Cl. Connected perimeters C2. Radiating interior
with reserved wetlands development
(incomplete) (incomplete)

Figure 8. Development patterns. Configurations of golf course development, from
higher quality in level A, less desirable in levels B and C. A2 is a schematic,
others are examples seen in the Collier County planning maps. Each plot is 1 section,
260 hectares. Clear tan areas are the golf courses and accompanying forest stands.
Green patches are reserved wetland areas. Other colors: white, to be developed
residential; brown, agricultural; pink and purple, commercial. Housing areas are
represented by subdivided lots and street patterns.





37

Demographics and Habitat Use


Radio-telemetry produced 2497 tracking points and accompanying records on 29

individuals at Site 1 between December 1995 and July 1997, and 254 tracking points at

Site 2 between January and July 1997. These data were used to examine population

structure, size and location of home ranges (Appendix C), nesting sites, habitat use, and

feeding patterns.

Survival

Eighteen fox squirrels were collared at Site 1 in the first year, 9 males and 9

females. In the second year, 11 more individuals were added to the study, 4 females and 7

males. Adult male fox squirrels had lower average survival rates than adult females (Table

2). Three individuals were known dead or moved to a neighboring course as adults. Four

individuals disappeared from the population and were not located again on the Cypress

course or on neighboring courses.

Adult females had an average 6 month survival of 0.87, the highest rate at Site 1.

Two adult females disappeared in the fall of 1997. One had a home range between 2

aggressive females and was regularly chased by each. It is possible this female moved west

into the Hole in the Wall and Wilderness area, though she was never seen on the courses.

The second female disappeared the same day her collar was found at the base of a tree

near a busy cart path.

At 0.78, the survival of subadult males was higher than adult males and subadult

females (Table 2). The immediate fate of all subadult males was known. In the first year of

study, 3 of the 4 remained on the course and moved into the adult cohort, and the












Table 2. Survival and birth rates for Site 1. Rates calculated in 6 month intervals. Survival equals persistence in the course
population. Birth rates are calculated from the number of young emerging from the nest. Weighted averages for each sex
and age class are the sum of persistent individuals in the 3 time periods divided by the sum of n for the 3 time periods.

SITE 1 n Known Moved or Disappeared Persistent in Survival Young Births/ Births/
FEMA S dead in dispersed population rate from nest adult adult
FEMALES population female
adults
Jan.-June 1996 4 0 0 0 4 1.00 4 1.00 0.50
July-Dec. 1996 6 0 0 2 4 0.67 15 2.50 1.25
Jan.-June 1997 5 0 0 0 5 1.00 3 0.60 0.23
15 0 0 2 13 Weighted 22 1.47 0.67
Saverage
subadults
Jan.-June 1996 5 0 3 0 2 0.40 0
July-Dec. 1996 0 0 0 0 0 0
Jan.-June 1997 3 0 0 0 3 1.00 0
8 0 3 0 5 0.63 Weighted 0
I average

MALES-
adults
Jan.-June 1996 4 0 1 1 2 0.50 Average adult 6
July-Dec. 1996 6 0 1 0 5 0.83 0.76 month survival
Jan.-June 1997 8 1 0 3 5 0.63
18 1 2 4 12 067 Weighted Average subadult 6
1t 11 I - average 0..71 month survival
subadults
Jan.-June 19% 4 0 0 0 4 1.00 Average juvenile 6
July-Dec. 1996 1 0 1 0 0 0.67 month survival
Jan.-June 1997 4 1 0 0 3 0.75
9 1 1 0 7 Weighted
I 0. average





39

fourth dispersed to Bear's Paw course and later disappeared. In 1997, 3 of 4 subadult

males survived to the end of the study and stayed on the course. The fourth died while

infected with skin fungus (Dr. Sharon Taylor, personal communication).

Subadult females showed the greatest change in survival rates from one season to

another (Table 2). In winter/spring 1996, 3 of the 5 subadult females dispersed from the

course, for a 6 month survival rate of 0.40. Two of these were found dead on other

courses, 1 within the summer and the other within the year. The third dispersing female

disappeared from a developed neighboring course. In winter/spring 1997, the 3 collared

subadult females remained on the course in their natal home ranges though the end of the

study. At that time they ranged in age from 11 to 13 months. None had reproduced

though 1 was known to be the object of a mating chase.

Trapping difficulties in the summer 1996 meant than no subadults from winter

litters could be collared. All subadults collared in the winter of 1995-1996 dispersed by

August, or they entered the adult cohort.

The survival data for Royal Palm, Site 2, covered a shorter period of time, only 7

months (Table 3). Four adults were collared, 2 males and 2 females. All remained alive

throughout the 7 months. Four subadults were collared, 2 male and 2 females. None

remained on the course at the end of 6 months. One male disappeared shortly after

collaring. Both females died: 1 following a severe infestation of skin fungus, the other

after being hit by a vehicle on Augusta Boulevard. The surviving subadult male moved

from the course through a series of progressively more distant feeding sites. At the end of

the study he was living near a feeder on the edge of Hibiscus course along Highway 41.







40




Table 3. Survival rates for Site 2, Royal Palm, for January 1997- July 1997.
Survival equals persistence in the course population. No births, b(x), were recorded
in the collared population between January and July 1997.

n Known Moved/ Disappeared Persistent in Survival/6
SITE 2
SE2 dead dispersed the mo
MALES- population
Adult 2 0 0 0 2 1.00
Subadult 2 0 1 1 0 0.00

FEMALES b(x)
Adult 2 0 0 0 2 0 1.00
Subadult 2 2 0 0 0 0 0.00






41


Reproduction

Reproduction at Site 1 varied widely between seasons and between individuals

(Table 2). Winter reproduction was lower in 1995-1996 and 1996-1997 than the

summer/fall of 1996, with only 4 and 3 young known to leave the nest in the 2 cooler

seasons. The 6 month rate of reproduction for the first winter season was 1.00/adult

female, in the second winter season it was 0.60/adult female. Winter young were born in

December through February, emerging from the nest in January through April. The

summer of 1996 was one of high reproduction. Five of the 6 adult females produced

evident young, with 2-4 young seen emerging from each of the 5 nests, for a total known

reproductive output of 15, a rate of 2.50/adult female for the 6 months. Warm season

young were born from early July to September and emerged form the nests between

August and late October. Mating chases were recorded in April though July and in

October.

Neither female at Site 2, Royal Palm, produced young from a nest during the

course of the study. One female was obviously pregnant at the time of collaring and

appeared to tend a brood nest for a few weeks. She failed to show signs of long-term

nursing and no young emerged from the nest. She appeared to be tending a brood nest

when her collar was removed, as did the other female in the study. The surviving subadult

male in the study was the offspring of one of the collared adult females, undoubtedly born

in the summer of 1996. At the time of his collaring, he and another subadult regularly

accompanied her. Though reproduction was not observed in collared animals during the

study, some adult females were successfully reproducing at Site 2, though apparently at a

lower rate than Site 1.






42


Mortality and Disease

Known causes of death in the collared populations of Sites 1 and 2 were:

automobiles (2), skin fungus infection (2), and electrocution (1). Three subadult

uncollared fox squirrels at Site 1 also died from vehicle accidents, 1 from a car on the

entrance road and 2 from golf carts on the course. Of the 5 squirrels known to be killed by

cars or carts at the study sites, only 1 remained at the site of impact. The others moved

away, sometimes several meters, before they died.

Skin fungus (Dr. Sharon Taylor, person, commun.), causing heavy fur loss and

blackened crusting of the skin, was apparent in both populations in 1997, affecting at least

8 collared individuals. One subadult died at each site. It primarily affected subadults,

though also was seen in 2 adults. The proportion of affected individuals appeared similar

in collared and uncollared squirrels. One uncollared subadult died at Site 1 within a month

of the collared individual. Fur loss and darkened skin as seen in a skin fungus infection was

observed in squirrels at courses adjacent to the study sites prior to 1997. It was easily

transmitted by contact and feeding areas appeared to be vectors, especially when squirrels

fed from a concentrated food supply. At least 3 squirrels with severe fur loss and skin

damage were seen sharing a feeder at CCN. The collared individuals who survived the

spring 1997 skin fungus infestation regained a thick, healthy coat in the late spring molt.

Information on predation is slight. A bobcat was seen killing a ground feeding,

uncollared fox squirrel at Site 1 in 1997 (A. Grieser, person. commun.). At least 1 female

bobcat and 2 young were regularly seen in the forest areas adjacent to Poinciana on the

east and west. The adult bobcat began frequenting Poinciana in daylight hours when the

area immediately east of the course was being cleared for development in 1997. There is






43

no evidence of raptor kills on the study sites, though I did find apparent raptor-killed fox

squirrels at 2 other nearby courses.

Population estimates

Estimates of population size, derived from repeat squirrel counts at the study sites,

are presented in Table 4. Range of minimum number alive (MNA) at Site 1 is 15-26. The

Lincoln estimates range from 15.9-32.5 and the Bailey estimate for small samples is 15.6-

30.5. Counts in July were understandably lower as a number of subadults had dispersed

from the course and young of the season would not emerge from nests until early fall. At

Site 2, the MNA range is 3-8, the Lincoln estimates range from 8.3 to 15, and the Bailey

unbiased estimates are lower at 7.5 to 11.7.

RAMAS Ecolab software (2.0, Applied Biomathematics) calculated an

annual growth rate (X) of 0.803 using the Site 1 winter/spring 1997 reproductive rate of

0.23 young/adult for 6 months, lowest of the 3 periods. Use of the summer/fall 1996

reproductive rate of 1.25 young/adult gave an annual growth rate of 1.39. The average

birth rate/adult for the 3 periods of study, 0.67, yielded an annual growth rate of 1.10.

Home range

At Site 1, in the first year of study, adult female home range size varied from 8.0 to

12.0 ha (x=10.1 ha) (Table 5). Home ranges of the 6 adult females cover the study site

(Fig. 9) from the north end to the southern border. No other adult females were observed

to make exclusive use of the course during the first year of study. A light color female

occasionally fed in the northeast section of the study site, though the core area of








44






Table 4. Population estimates of Sites 1 and 2. Estimates are derived from repeat censusing of
each site.
Total Known Uncollared Collars MNA Lincoln Bailey Standard
seen collars in sample Index unbiased error of
Index Bailey
SITE 1 Index
14-Apr-96 19 16 8 11 24 27.6 26.7 4.68
28-Apr-96 7 15 3 4 18 26.3 24.0 6.00
6-May-96 18 14 5 13 19 19.4 19.0 2.52
25-May-96 10 13 6 4 19 32.5 28.6 8.62
6-Jun-96 11 13 5 6 18 23.8 22.3 5.09
1-Jul-96 9 13 3 6 16 19.5 18.6 3.60
1-Jul-96 11 13 2 9 15 15.9 15.6 1.92
1-Mar-97 14 19 5 9 24 29.6 28.5 4.96
20-Mar-97 8 18 2 6 20 24.0 23.1 3.86
29-Mar-97 19 18 7 12 25 28.5 27.7 4.38
6-Apr-97 15 17 7 8 24 31.9 30.2 6.32
21-Apr-97 16 17 7 9 24 30.2 28.9 5.59
11-Jun-97 20 16 10 10 26 32.0 30.5 6.08
Means 13.6 20.9 26.2 24.9

SITE 2
11-Jan-97 8 0 8 0 8
19-Feb-97 6 5 4 2 9 15.0 11.7 4.41
8-Apr-97 3 7 1 2 8 10.5 9.3 2.33
15-May-97 4 5 2 2 7 10.0 8.3 2.64
14-Jun-97 5 5 2 3 7 8.3 7.5 1.94
Means 7.8 11.0 9.2










Table 5. Home range summary by site and year. Home range size equals the 95% area (ha) as calculated by CALHOME, using kernel analysis.
Bold numbers are the mean of each series.

Adult females Subadults Adult Males
(ha) (ha) (ha)



Site 1, year 1, Dec. 1995 -Oct.31, 1996 8.0 10.7 42.2
8.8 14.6 46.0
10.1 15.9 76.5
10.7 20.2 118.4
10.9 22.6 70.8
12.0 49.1
10.1 22.2

Site 1, year 2, Nov. 1996-July 29, 1997 9.1 5.9 44.1
13.1 10.3 48.4
19.2 12.1 99.9
19.7 16.6 114.1
20.9 17.3 118.0
16.4 20.8 121.0
21.5 90.9
14.9

Site 2, year 2, Jan. 1997-July 29, 1997 13.1 25.8 136.1
30.6 40.8 303.8
21.8 108.5 220.0
58.3






46
































001, 10.9 ha
O 02, 10.2 ha
S03, 12.0 ha
05, 8.8 ha
S06, 10.7 ha
D 15, 7.9 ha

KM


Figure 9. Adult females, Site 1, year 1. Tracking locations and home range boundaries,
defined by the 95% contour, kernel analysis, CALHOME. The summer/fall brood nest
site of each female is identified by placement of her individual number.






47


her home range was east of the study site in the Pines course. Of the 431 tracking

locations recorded for the females between December 1995 and November 1996, only 10

were outside of the course boundaries, all but 2 of these 10 remained within the

boundaries of the Poinciana. Females did not range into neighboring pine or cypress stands

and only ROPO 6 crossed the entrance roadway on the club grounds to feed in a large

ficus immediately north of the road. All adult females showed some home range overlap

with other adult females, though none showed range overlap with more than 2 other adult

females.

In summer/fall 1996 all females maintained a brood nest in a location within a core

area outside the home ranges of all other adult females. In the first few weeks the of

summer brood nest occupancy females stayed in the nest most of the day and greatly

reduced the area in which they fed (Table 6). Placement of the nest within a small mixed

stand often allowed them to feed without moving to the ground. A similar pattern of

reduced movement and isolation was not observed during the winter brood nesting period.

Adult males at Site 1, year 1, had home ranges of 42.52 ha to 118.40 ha

(x = 70.84 ha) (Table 5), significantly larger than those of the adult females (Mann-

Whitney U=0, p<0.05). The home ranges of all 4 adult males overlapped in the center of

the study site (Fig 10). Other adult males were regularly seen on the course. ROPO 04 and

ROPO 07 used the Cypress course almost exclusively, while ROPO 16 used all of the

Cypress course and the adjoining back nine of the Pines course on the east. ROPO 17 used

portions of 4 courses, the Cypress, Pines, Hole in the Wall, and the Country Club of

Naples to the west. Though ROPO 17 used the open edges of large forested stands within






48





Table 6. Brood nest home ranges. Home range size for 3 adult females
occupying brood nests in summer/fall 1996, Site 1. Home range determined
by kernel analysis, CALHOME.

Brood nest home Brood nest core Number

range, 95% contour area, 50% contour sightings

ROPO 02 0.8 ha 16.7 m2 16

ROPO 03 0.3 ha 9.9 m2 13

ROPO 06 2.1 ha 78.5 m2 9







49



































S04,46.0 ha
07, 42.5 ha
0 16, 76.5 ha
1317,118.4 ha 4



KM
.5

Figure 10. Adult males, Site 1, year 1. Tracking locations and home range boundaries,
defined by the 95% contour, kernel analysis, CALHOME.





50


and between courses, he was never found in the interior of these stands. He did not need

to cross the roadway to move from CCN to HIW and was regularly observed moving to

HIW by crossing the canal between the Cypress course and HIW by way of a natural tree

bridge on the west side of fairway 2. These tree routes were frequently used by several

individuals in moving between these 6 neighboring courses separated by canals.

Home ranges of the 6 collared subadults at Site 1, year 1 (Table 5, Fig. 11), ranged

from 10.66 ha to 49.07 ha (x= 22.16 ha). ROPO 13, a male, began making long day trips

to the back nine of the Poinciana Pines course in the month prior to his dispersal. His large

home range, twice as large as the mean and the next largest subadult home range, reflects

these trips. The 4 subadults in the southern half of the study site show a strong overlap

(Fig. 12), with all four commonly using forested areas along the 7t and 13" fairways.

Three of these 4 dispersed to other courses between March and August 1996. The fourth

stayed on the Cypress course until the end of the study.

ROPO 10, a subadult female, dispersed from the southern section of Site 1 to

Poinciana Pines at the end of March 1996 and in early April moved to the CCN (Fig. 13 ).

Her initial movement to the front nine of the Pines course was 1.4 km with an additional

0.85 km to CCN. She also frequented a home feeder at point B and crossed Burning Tree

Drive regularly. She slipped her collar in June 1996 and was seen only once again, despite

subsequent searches.

ROPO 14, a subadult female, dispersed from the northeast section of Site I in late

April 1996 (Fig. 14), moving to Bear's Paw Country Club. Her dispersal distance from her

Site 1 to her first sighting at BP was 2.8 km. She occasionally fed on scraps put out at Site

A, with the remaining tracking sites at the edges of heavy forest on the northeast edge






51
















































prior to dispersal.
008, 14.6 ha
109, 22.6 ha *
O 10, 20.2 ha *
3 12, 15.9 ha
13,49.1 ha*
S314, 10.7 ha *


F - .5


Figure 11. Subadults, male and female, Site 1, year 1. Home range boundaries are
defined by the 95% contour, kernel analysis, CALHOME. Individuals that dispersed
during the 1996 spring and summer seasons are marked (*), home ranges are those
prior to dispersal.






52

































S 08, 14.6 ha
S09, 22.6 ha *
310, 20.2 ha *
13,49.1 ha *




tKM


Figure 12. Four subadults, Site 1, year 1.Collared subadults in the southern half of the
course. Tracking locations and home range boundaries indicate use of the course.
Three of 4 squirrels, 2 females, 1 male, dispersed from the course in spring and
summer 1996. Only ROPO 08 remained through year 2 of the study.







53















































KM
.5

Figure 13. Dispersal movement of ROPO 10. Movement from Site 1 to Country Club
of Naples in late March, 1996. Movement from her central home range area to A was
1.4 km, and from A to B an additional 0.85 km, for a total 2-week dispersal distance
of 2.2 km.







54






































M





GOLDE GATE AKWA )







I I

Figure 14. Dispersal movement ofROPO 14. Dispersal in April 1996 to Bear's Paw
Country Club. The distance from her central home range area to the first off-site
location, A, was 2.8 km. She was found dead at B in late July 1996.






55

of the course. In July she was limping badly (L. Molden, person, commun.) and was

found dead in dense undergrowth on July 24, 1996. At least 1 large cat regularly roamed

the small area where she was found.

In mid-May 1996, ROPO 09, a subadult female, moved from the center of Site 1

to the eastern end of the Quail Run Golf Course (Fig. 15), a heavily developed course with

few tree stands, a dense street network and heavy traffic. By the end of June 1996, ROPO

09 had moved to the CCN where she often fed at a home feeder station used by 2 or 3

other fox squirrels, at least 2 of which had severe fur loss with accompanying thick,

darkened skin, probably related to a skin fungus. By August, ROPO 09 was suffering from

a similar fur loss when she was tracked to the center of the HIW. She returned to Site 1,

Cypress course, for a brief period in September 1997. Regular contact was lost in early fall

due to collar failure. She continued to use the HIW and the south end of the CCN, where

she frequently crossed roadways in her daily movements. She was found dead at the side

of the entrance road to CNN in April 1997. Her initial dispersal distance was 2.6 km and

the total distance moved from the original home range to HIW was 6.0 km.

ROPO 13, a subadult male, dispersed from the southern section of Site 1 to Bear's

Paw Country Club in August 1996 (Fig. 16). On August 8, 1996, he moved from the

south border of Site 1 to Golden Gate Parkway, through 1.2 km of pine forest. The

following morning he crossed the busy 4 lane roadway into BP, where he was tracked

until mid August 1996. His initial 24 hour dispersal distance was 1.9 kmn his total

movement distance from his original home range was 3.5 km.






56






A

































KM


Figure 15. Dispersal movement of ROPO 09, mid May 1996. First off-site location at
A, a move of 2.3 km from previous home range on May 20, 1996. Located at B on
May 27, 1996, at C, home feeding site on June 28, and at D on August 23. She was
found dead at E, hit by vehicle, in April 1997.






57





























CID,














Figure 16. Dispersal of ROPO 13. Movement from Site 1 to Bear's Paw Country
Club on August 8, 1996. Points north of A are tracking locations of 13 before that
date. Movement from A to B, 1.2 km, took 3 hours. He crossed from B to C the
following morning, for a total 24-hour dispersal distance of 1.5 km. The total
distance traveled to his final sighting was 3.5 km.







58



































0 02, 13.1 ha
r dl 03, 192 bha
05, 20.9 ha
3 15, 19.7 ha
S28, 9.1 ha


KM
.2

Figure 17. Adult females, Sitel, year 2. Tracking locations and home range boundaries,
defined by the 95% contour, kernel analysis, CALHOME. Numbered site is 1997
summer brood nest of female 03.





59


At Site 1, year 2, the home ranges of the 5 adult females varied from 9.08 ha to

20.92 ha (x=16.40) (Fig. 17). The 4 adult females collared from the previous year,

showed significant increases in home range size (Table 5) (t=3.95, P=0.029, df=3) though

each female continued to use approximately the same section of the course. ROPO 03 and

ROPO 05 increased use of the edges of large forest stands on the eastern edge of HIW

and ROPO 15 used open edges of forested roughs at Wilderness Country Club and a

feeder area on private property adjoining the southeast corer of Site 1. ROPO 03

expanded to the north into the previous home range of ROPO 06, who disappeared in

December 1996. Core areas of ROPO 03, ROPO 05, and ROPO 15 showed increases

from year 1 to year 2. ROPO 15 and ROPO 28 had high overlap, though location of core

areas indicated that ROPO 28 made heaviest use of the southeast portion of the course

and ROPO 15 the southwest corer of the corer.

Only ROPO 03 was tending a summer brood nest when the study ended in July

1997. The second year home range of ROPO 05, a female, had expanded to include the

fall 1996 brood nest site of ROPO 03. ROPO 03 moved her 1997 summer brood nest to

the north, into a tree stand formerly within the home range of ROPO 06, an adult female.

The 6 adult males of Site 1, year 2, used portions of 5 courses (Figs. 18, 19).

Home range size varied from 44.06 ha to 121.00 ha (x= 90.91 ha) (Table 5), significantly

larger than adult female home ranges in the second year (Mann-Whitney U=0, p<0.05).

All of the males used at least some part of 2 or more courses and did so with little crossing

of roadways. ROPO 20 used much of the Cypress course, visited the front and back

sections of the Pines course and frequently moved into Hole in the Wall. ROPO 18 rarely

visited the Cypress course, spending most of his time in the northern






60






































bo 08, 44.1 ha
020,121.0o ha

22, 48.4 ha


KM


Figure 18. Three adult males, Site 1, year 2. Tracking locations and home range
boundaries, defined by the 95% contour, kernel analysis, CALHOME.







61








































S07, 114.1 ha
11, 99.9 ha

0 18, ll8.0ha


KM
.5

Figure 19. Three adult males, Site 1, year 2. Tracking locations and home range
boundaries defined by the 95% contour, kernel analysis, CALHOME.






62


section of the Pines course and the CCN. ROPO 07 and ROPO 08 used the Cypress

course and both sections of the Pines course. ROPO 08 and ROPO 20 had the smallest

home ranges, each using portions of the Cypress course and the back nine of the Pines

course. Males again showed strong overlap of home ranges.

Seven subadults at Site 1, year 2, had home ranges of 5.88 ha to 21.47 ha

(x=14.93) (Figs. 20 & 21). All subadults show overlap with at least 3 other subadults.

None of the subadults dispersed before the end of tracking in July 1997, though ROPO

21, a male, began to use the northeast corner of Wilderness Country Club to feed in June

and July 1997.

Five of the subadults at Site 1, year 2, were born to adult females ROPO 02,

ROPO 03, and ROPO 05 who remained on the course in the same home range areas

throughout the study. Figures 22 and 23 compare the 1997 spring/summer home ranges of

the collared offspring to the 1997 home ranges of their mothers. The female offspring of

ROPO 02 and ROPO 03 had home ranges entirely contained within those of their mothers,

while the male offspring of ROPO 03 and ROPO 05 had home ranges that extended

beyond those of their mothers. The overlap of the subadult home ranges with the core area

of their mother, ROPO 05, is clear (Fig. 23).

Adult female ROPO 01 disappeared in December 1996 and offers no comparison

with her 1997 home range area with those of her offspring, ROPO 21 and ROPO 24.

Nevertheless, it is clear that her 1996 offspring, male ROPO 21 and female ROPO 24,

continued to use their natal home range, the core area of female ROPO 01 before her

disappearance (Fig. 24). ROPO 24 expanded to the northeast in her mothers absence and

ROPO 21 expanded to the south and west.






63



































25, 12.11 ha

1326, 20.8 ha
I 29, 1.3 ha






Figure 20. Three of 7 collared subadults, Site 1, year 2. Tracking locations and home
range boundaries as defined by the 95% contour, kernel analysis, CALHOME.







64






































O 21, 16.6ha


O 24,21.5ha
27,5.9 ha



.2


Figure 21. Four of 7 collared subadults, Site 1, year 2. Tracking locations and home
range boundaries as defined by the 95% contour, kernel analysis, CALHOME.







65


































-26,20.8 ha
27, 5.9 ha
29, 10.3 ha
303,19.2 ha
02, 13.1 ha



L --- M-------

Figure 22. Two adult females and offspring, Site 1, year 2. Females, 03 and 02, and their
collared offspring from summer 1996. Thin lines are subadult offspring. Squirrels 27
and 29 are females, 26 is male.







66




































O'] 23, 17.30 ha

S 25, 12.11 ha

05, 20.9 ha


(KM
I I


Figure 23. Home ranges of adult female and offspring, Site 1, year 2. Adult female 05
and collared male offspring 23 and 25 from summer 1996. Yellow dots indicate the
core area, 50% contour, of female 05.







67
































SH r i, 21, 16.6 ha
1 24, 21.5 ha
0 01 oo
[Z21 core

024 core

KM


Figure 24. Home ranges of female and 2 offspring, Site 1, year 2. Subadults 21 and
24, siblings from female 01 in the summer 1996, show overlap of home ranges and
core areas. Core area of 01, who disappeared in December 1996, is shown in light
blue.





68

All 7 of the Site 1, Year 2, subadults continued to use their natal homeranges for

the first year of their lives, 5 sharing with their mothers, 2 remaining after their mother

disappeared.

Seven of the 8 squirrels collared at Site 2 in year 2 remained to provide usable

home range data. ROPA 01 and ROPA 08, adult males, had home ranges of 136.1 ha and

303.8 ha, respectively (Table 5). ROPA 01 used most of Site 2, Royal Palm, while ROPA

08 used Site 2 and most of the neighboring Hibiscus Country Club (Fig. 25). ROPA 08

readily moved from the east to the west end of his home range, a distance of 2.5 km,

within 24 hours. Both males regularly crossed Augusta Boulevard, while their movements

on either side of that busy street generally followed the fairways and appeared to minimize

travel through housing. Though their home ranges overlapped, they were never seen

together as males at Site 1 often were.

The two adult females at Site 2 had home ranges of 13.06 ha and 30.57 ha (Table

5, Fig. 26). Adult female ROPA 04 had a home range 50% larger than any at Site 1. She

often crossed Augusta Boulevard. Adult female ROPA 06 had a home range similar in size

to those at Site 1. Her home range included a regularly stocked feeder at a private

residence. On rare occasions she crossed into the central pine stand within the private

housing area. Though the home ranges of these 2 adult females were widely separated, no

other adult females where ever observed in the area between their 2 home ranges.

Three of the 4 collared subadults at Site 2 provided data on home range size, these

varied from 25.77 ha to 108.50 ha (x= 58.34 ha) (Table 5, Fig. 27), significantly larger

than Site 1 subadults for the same period (Mann-Whitney U=0, p<0.05). Male ROPA 2










L 01, 136.1 ha
O08,303.8 ha
* o points
08 points
Feeders



















KM

Figure 25. Adult males, Site 2, 1997. Tracking locations and home range boundaries as defined by the 95% contour,
kernel analysis, CALHOME. Yellow crosses mark the locations of feeders.







70




































3 04,30.6 ha
306, 13.1 ha
S04 points
S06 points



.2

Figure 26. Adult females, Site 2, 1997. Tracking locations and home range boundaries
as defined by the 95% contour, kernel analysis, CALHOME






























03, 40.8 ha
05, 25.8 ha
07, 108.5 ha
S02
* 03
S05
07



Figure 27. Subadults, Site 2, 1997. Tracking locations and home range boundaries as defined by the 95% contour, kernel analysis,
CALHOME






72


was an older subadult who disappeared within a month of being collared and provides no

usable home range. Female ROPA 3 used an area on both sides of Augusta Boulevard and

across Forest Hills Boulevard into the Hibiscus course. Female ROPA 05 used an area on

both sides of Palmetto Dunes Circle. Male ROPA 07, summer 1996 offspring of adult

female ROPA 06, had an unusual home range use pattern, sequential use of small patches.

He spent a few weeks to a month in a small area of a hectare or less and then moved to

another area, each time moving away from his natal home range at the northwest corer of

Site 2. In the final week before his collar was removed he had moved to the west side of

Hibiscus Country Club along U. S. 41, 1.2 km from his natal home range.

Nest Sites

At Site 1 fox squirrels made regular use of untrimmed or lightly trimmed cabbage

palms, bromeliads and cypress cavities for sleeping nests. Stick nests were used on

occasion. The common use of untrimmed palms and bromeliads as nests eliminated the

possibility of counting nests and determining the nest to squirrel ratio. Squirrels were

regularly observed carrying Spanish moss (Tillandsia usneoides) to nesting sites. Squirrels

rarely constructed open platform nests for daytime resting but simply draped themselves

along branches.

Brood nests were readily observed at Site 1 in summer 1996 as females seldom

moved from the nests for 2 -3 weeks. Nests were located in mixed stands of pine,

untrimmmed or lightly trimmed palms, and cypress (Fig. 9). Three females used cavities

high in large cypress trees, 1 raised a litter of 4 from such a cavity, another a litter of 3.

Two females raised young in the center of densely leafed palms, one a queen palm

(Arecastrum romanzoffianum) and the other a cabbage palm. The remaining female used a






73


large bromeliad high in a large pine for her nest site. The nest was located in the base of

the large plant. All females used Spanish moss in constructing and maintaining nests. The

queen palm nest also contained shredded queen palm leaflets and required her to spend

time stripping leaflets and tearing them into strips. Winter brood nests were similar to

those in the summer of 1996 and included 2 in cypress cavities, 2 bromeliad nests, and 1

palm nest.

As the study ended at Site 2, both adult females appeared to be tending brood

nests. One was in a rather isolated moderately trimmed cabbage palm near a canal and the

other was in a stick nest in a moderate-sized pine tree. In the winter of 1997, the same

female was observed using a wood duck box with her offspring from the previous

summer. All 3 nested together in the box during times of heavy rain, cooler nights, or high

wind.

Landscape Composition and Vegetation Mapping

Landscape composition of Sites 1 and 2 and the Poinciana Pines course is

presented in Table 7. While Site 2 was 2.3 times larger than Site 1, the golf course and

tree stands within the roughs of the 2 courses were similar in size, though not necessarily

similar in species composition or structure. The obvious difference in landscape

composition between the 2 sites was the presence of housing areas at Site 2. Twenty-eight

percent of Site 2 was occupied by residential development, streets and clubhouse property,

and an additional 25% was condominium land and a private pine stand within a housing

area. The 2 courses that comprised Royal Poinciana Golf Club, Site 1 and the adjoining

Pines course, were similar to one another in general landscape composition. The smaller






74



Table 7. Landscape composition. Landscape cover of Site 1, Site 2, and Royal Poinciana
Pines course.
Royal Poinciana Royal Palm Royal Poinciana
Cypress Site 2 Pines Course
Site 1
Area, ha % of Area, ha % of Area, ha % of
total total total
Total size of site 61.4 141.9 62.9
Club property in 30.0 49% 26.0 18% 24.8 39%
tree stands
Condominium land 0.0 0 34.9 25% 0.0 0
in trees
Residential, streets 0.0 0 40.1 28% 0.0 0
and clubhouse
Lakes, canals, 5.5 9% 8.0 6% 5.3 8%
wetlands
Fairways, greens, 25.9 42% 32.9 23% 32.8 52%
sandtraps,
driving ranges,
unforested
roughs






75


area of tree stands in the Pines course reflected the open nature of the back 9 of that

course and the presence of 2 driving ranges within the course.

While Site 1, Site 2, and the Pines course each had between 30.0 and 24.8 ha of

tree stands or forested area, the species composition and structure of these plots were not

alike. Site 1 sampling plots contained a greater diversity of tree species than Site 2 plots.

(Table 8, Fig. 28,29). A mixture of native tree species, pine, cabbage palm, and cypress,

dominated Site 1. Though Site 1 was high in native hardwoods, there were no class 1, pine

dominant, stands. Pines were found as co-dominants with cabbage palm (11%), cypress

(3%), or both (10%). Cypress was dominant in 14% of the forested area and was co-

dominate with pines and/or cabbage palms in an additional 31% of the area. Palm

dominant stands accounted for 8% of the area in tree stands. At Site 1, 64% of the tree

stands were dominated by the pines, cypress and cabbage palms. A mixture of native

species, often including oaks (Quercus virginianum, Q. laurifolia) , maple (Acer rubrum),

red bay (Persea borbonia) made up another 10% of the forested landscape. While native

species dominated the landscape, the importance of exotic species on the Cypress course

was seen in the extent of class 9 (mixed natives with exotics), which covered 26% of the

forested area.

Site 2 was dominated by pine, with 68% of the plot area in class l(pine dominant)

and an additional 6% with pines as co-dominants (Table 8, Fig. 29). Cypress was a minor

element of the Site 2 vegetation, dominating in only 2% of the area. Classes 6 and 7 of the

mixed natives were not present at Site 2. Exotics were less common than at Site 1, with

class 9 (mixed natives with exotics) accounting for 11% of the area.






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Table 8. Vegetation classification of Site 1, Site 2, and Royal Poinciana Pines.


Royal Royal Palm Royal
Poinciana Site 2 Poinciana
Cypress Pines Course
Site 1
Vegetation classes, by relative Area, % Area, % Area, %
basal area ha ha ha

1-Pine, 70% or more pine 0.0 0 17.8 68 3.0 12
2-Pine and cabbage palm, 3.2 11 0.9 3 2.4 10
30% or more of each
3-Palm, 60% or more cabbage 2.4 8 1.4 5 2.0 8
palm
4-Cypress, 60% or more 4.4 14 0.4 2 3.5 14
cypress
5-Pine and cypress, 30% or 0.9 3 0.7 3 0.0 0
more each of pine and
cypress
6-Pine, cypress, c. palm, over 2.9 10 0.0 0 0.0 0
85% of the 3 combined,
with each being over 20%
7-Cypress and c. palm, 30% 5.5 18 0.0 0 0.0 0
or more of each cypress
and pine
8-Mixed natives-none of the 2.9 10 2.1 8 1.2 5
above with 20% or less
exotics
9-Mixed natives with exotics- 7.8 26 2.8 11 12.7 51
not 1-7, with more than
20% exotics






77




























Vegetation Classes
1-Pines
-Pine & cabbage palm
3-Cabbage palm
4-Cypress
5 -Pine & cypress
lPine, cypress & c. palm
7- Cypress & c. palm
S8-Mixed natives
9-Mixed with exotics





Figure 28. Tree stand characteristics at Site 1, Royal Poinciana Cypress. Nine
vegetation classes are color coded. H= high density (>200 stems/ha), L=low density
(<100stems/ha), all patches not marked H or L are moderate density (100-199 stems/ha).
Presence of dense litter layer is indicated by an asterick (*); S= presence of a shrub
layer. Light gray areas between tree stands are the fairways, tees, and greens.


























Vegetation ses 00
I -Pine
S 2-Pine & cabbage palm
3-Cabbage palm
4-Cypress
5-Pine & cypress
6-Pine, cypress& c.palm
7-Cypress & c. palm
8-Mixed natives
9-Mixed with exotics a M
.5

Figure 29. Tree stand characteristics at Site 2, Royal Palm. Nine classes are color coded. H--high density (>200 stems/ha), L=low
density (<100 stems/ha),VL=Very Low density (<30 stems/ha). All patches not marked are moderate density (100-199 stems/ha).
Presence of litter layer is indicated by an asterisk (*). Areas along Augusta Boulevard that are outlined in dark green and contain
letters indicating tree density are the condominium properties. The outline area in the upper left is the private pine forest.






79


























Vegetation Clasres
1-Pine
2-Pine & cabbage
S3-Cabbage
4-Cyp rss
D 5-Pine &cypress
S 6-Pine, cyess & c.a an a p n a h a a
S -Cypreess & . palm
8-Mixed natives
9-Mixed with exotics


KM

Figure 30. Tree stand characteristics for Royal Poinciana Pines course. Nine classes are
color coded. H=high density (>200 stems/ha), L=low density (<100 stems/ha), all
patches not marked H or L are moderate density (100-199 stems/ha). Presence of a
dense litter layer is indicated by an asterisk (*); S= presence of a shrub layer. Gray
areas between tree sands are fairways.






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The Pines course had a high presence of exotic species, with 51% of the tree stand area in

class 9, and a much lower area dominated by a mixture of pines, cypress and cabbage

palms (Table 8, Fig. 30). Pine dominated stands (class 1) accounted for 12% of the

forested area and pine/cabbage palm for an additional 10%. Cypress dominated 14% of

the area. Classes 5, 6 and 7, mixtures of pine cypress and cabbage palm, were absent from

the Pines course.

Structure of the trees stands on the 3 courses also differed from course to course

(Table 9, Figs. 28-30). Site 1 had a higher density of trees than Site 2, with 24.9 ha of

forested area having more than 100 stems/ha and only 5.2 ha in lower density stands. Site

2 had a much more open landscape, with 17.9 ha of low density (< 100 stems/ha) and only

8.4 ha in high and medium levels. The Pines course was a mixed course, with a dense

forested front 9 on the north and an open back 9 on the south.

The presence of a litter ground cover, as opposed to grass or bare soil, and the

presence of a shrub layer are noted in Figures 28-30. Site 2 had 4 plots with a significant

litter layer. All were in pine dominated stands, only 1 with a high tree density. Site 1 had 8

plots with heavy litter layers, 5 of these plots had high tree density and all of these stands

were dominated by a mixture of native pine, cypress or cabbage palms. Four plots at the

south end of Site 1 had a shrub layer, with wax myrtle (Myrica cerifera) the most

common understory species. The Pines course had 7 plots with a litter layer and 5 of these

had high tree density. The more open back nine of the Pines contained 2 large plots with

heavy litter layers and high to medium density of trees.

The condominium areas and private forest stand at Site 2 were dominated by

pines, though tree density varied widely with the number of buildings and parking lots in






81



Table 9. Density of trees at Site 1, Site 2, and Royal Poinciana Pines.

High Medium Low
> 200 stems/ha 100-199 <100 stems/ha
stems/ha
Site 1, R. P. Cypress-
area in ha 5.4 19.5 5.2
% 18% 65% 17%
Site 2, Royal Palm
area in ha 0.7 7.4 17.9
% 3% 28% 69%
Royal Poinciana Pines
area in ha 7.8 9.9 7.1
% 32% 40% 28%





82


each plot (Fig. 29). Most condominium plots had low pine densities, with 19.8 ha in the

range of 30-99 stems/ha and 5.6 ha at < 30 stems/ha. High density pine forests (>200

stems/ha) were found in 4 plots totaling 4.5 ha and 2 plots totaling 4.0 ha had moderate

densities (100-200 stems/ha). The 6 undeveloped private home vacant lots bordering the

course in the summer of 1997 had dense stands of pine with scattered cabbage palms. The

developed lots that circled the course on all but Augusta Boulevard had a very low density

of trees, with 3 or fewer trees in most lots. Pine and queen palms (Arecastrum

romanzoffianum) were the most common species. Black olive (Bucida buceras), cabbage

palms, oaks and bottle brush (Callistemon rigidus) were scattered throughout the lots of

private residences.

Habitat Use

Locations of 2138 of the 2497 tracking points of the Site 1 collared population

were used to examine habitat preference on the Royal Poinciana Cypress and Pines

courses. The 359 points not used were located either on neighboring courses, fairways or

unforested roughs. Comparison of the aggregated area of all plots in each of the 9

vegetation classes and the number of points located within the boundaries of each class

shows a non-random use of forested areas (Table 10, Fig. 31). On the Cypress course, all

subsets of the population showed non-random use of the forested stands. In all cases, fox

squirrels had higher than expected use of class 2 plots pine/cabbage palm co-dominated

stands While class 2 plots were 11% of the forested area, they accounted for 18% of the

tracking locations for females and 28% for males. A preference for class 2 plots was

shown in both years of the study.






83


Table 10. Habitat preference tests on tracking points at Royal Poinciana
Cypress and Pines courses. Chi-square tests the hypothesis that points are
randomly distributed among vegetation types. Area of each vegetation class
predicts the number of points. Only tracking locations within the mapped
plots of forested stands are included. Double asterisk indicates results are
significant at the 0.01 level.
Number of
Tracking X2 df p
Points
Cypress
Females, both years 1206 154.08 7 5.60E-30
Males, both years 768 242.76 7 9.60E-49
Year 1, all squirrels 1036 208.12 7 2.18E-41
Year 2, all squirrels 938 128.51 7 1.28E-24 *
Summer 1996 276 81.57 7 6.59E-15
Summer 1997 340 41.29 7 7.12E-07**

Pines
All points 164 20.73 5 0.0009**
Males, both years 140 17.59 5 0.0035*
Year 1 all squirrels 65 7.99 5 0.1570
Year 2 all squirrels 99 18.31 5 0.0026*
















20% _
* females,all
0 males,all
15% yr
Syr2
10% - U summeri









% -'-
6%









Figure 31. Difference in expected and observed use of vegetation classes at Site 1, Cypress course, by subgroups of the
tracked population.The nine classes of tree stands are shown on the x axis. Positive values indicate a stronger than
expected preference for a given vegetation class, negative values indicate less than expected use of a vegetation class.
Table 10 presents Chi-square results for tests of random distribution among vegetation classes for each of the subgroups
shown here.
shown here.





85


Site 1 fox squirrels showed consistent underuse of cabbage palm dominated plots,

cypress dominated stands and plots of mixed natives. Females showed a preference for

pine/cypress dominated plots, as well as pine/cypress/cabbage palm stands and

cypress/cabbage palm forested areas (Fig. 31). Males underused or showed expected use

of all vegetation types other than the pine/cabbage palm plots.

On the Pines course the aggregation of all tracking points within the tree stands

shows a preference for pine plots, mixed natives and natives with exotics. Both cabbage

palm plots and cypress stands were underused (Fig. 32). Females used the Pines course so

rarely, only 24 points in the 2 years of the study, that they were not examined separately

for patterns of use. The summer subsets were similarly small and not separately analyzed.

In the first year of the study, fox squirrel use of tree stands at the Pines course indicated

no significant departure from expected (Table 10).

The mixture of golf course property with private and condominium property at

Site 2 created a more complex landscape than seen at Site 1. Analysis of the 147 tracking

locations within the 26.0 hectares of tree stands on the Royal Palm property showed that

squirrels used the vegetation classes in proportion to their abundance on the landscape

(X2=12.22, df=6, p=0.058). Pine plots account for 68% of the forest stands and contained

65% of the analyzed tracking points. Class 9, mixed natives with exotics, and class 3,

cabbage palm dominated stands, had slightly higher than expected use. The 34.9 ha of

condominium property and the private pine forest inside the back 9 of the course had only

34 tracking points. Nineteen of these were in the 1.4 ha private pine forest that contained

at least 3, well-stocked feeders. The 40.1 ha of residential property accounted for only 17

tracking location, 10 at feeders.






86






60%

5% Expected
SAll Use
40%


2o%




0%
V a
0% IIIIj







Figure 32. Habitat preferences for all fox squirrels at the Pines course, Royal
Poinciana. All tracking points within the mapped vegetation plots on the Pines
course are included, n=164.











Table 11. Feeding patterns of fox squirrels on the 13 main food species eaten by the collared population of Site 1. Data
include all feeding records, at all locations used by the Site 1 collared squirrels for 19 months, n=817. Diversity Index
is the inverse Simpson measurement for richness and evenness. Cell shading :Section A, dark=250%, light=15-49%,
white=<15%, blank cells =0; Section B, dark=> 50%.
Section A- Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
Species data 96 97 96 97 96 97 96 97 96 97 96 97 96 97 96 96 96 96 96
%Natives
Pine 25 22 2 8 7 1 14 4 6 3 10 5 20C n i i 17
Cypress 42 37 18 4 5 2 1 6 4 16 21 21 20 23 18 19 33
Red maple 8 4 20 14
Oak sp. 7 6 3 8 4 10 13 30
Mushrooms 4 3 8 2 1 2 27 21 12 26 5 8 4
Larva 4 15

%Non-natives
Q. palm 8 7 33 22 7 5 16 4 4 18 3 6 16 5 8 6
Ficus sp. 4 20 2 6 16 4 3 17 39 11 23 21 5 4
Tallow 19 2 5 3 1 1
Bischofia 17 18 23 26 29 42 2 42 2 11 4
Bottlebrush 28 22 18 21 6 3 10
Silk oak 4 5 18 15
Java plum 3 12 35 4
n= 12 27 45 65 54 73 94 78 52 49 38 52 19 20 26 39 48 3 23


Section B-
Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
Summary
/Natives 75 70 27 49 4 23 5 24 15 41 39 58 53 45 88 92 94 100 100
%Non-natives 25 30 73 51 96 77 95 76 85 59 61 42 47 551 121 8 6 0 0

Diverst Index 3.6 4.3 4.4 6.0 2.5 6.11 3.2 5.3 4.1 3.8 3.8 6.2 5.1 4.5 2.1 2.7 2.51 1.8 2.5






88


Feeding Patterns

Feeding data from Site 1 show seasonal shifts between native and non-native food sources

(Table 11). Native foods, pine, cypress, and oaks, made up over 70% of the diet from

August 1996 through January 1997. In March through July of 1996 and 1997, non-natives

were the primary food source. February, June, and July of the 2 years showed a changing

mixture of native and non-native sources.

In 1996 squirrels began to feed on new, green pine cones in late June and early

July. By August cones were mature and pine became the dominant food, with squirrels

feeding alone or in small groups in or around select trees for several hours at a time. This

also began the season of burying cones, in grass, litter and sand traps. Squirrels were

observed burying in their core areas and at the edges of their home ranges. In December

and January, cypress cones increased in importance, as pines decreased. Cypress

constituted 20% or more of the diet for 7 months, July 1996 through January 1997.

Despite the tent caterpillar infestation of 1997, cypress were again accounting for 29%/ of

the diet in June and July of that year. Feeding on early pine cones began in June and July

1997.

Oaks were not a regular food source in 1996 until September, when oak feeding,

primarily live oak, accounted for 30% of the diet. Fox squirrels continued to feed lightly

on oaks through April 1997.

In both 1996 and 1997 fox squirrels took advantage of the bright red samara of the

red maples for concentrated feeding in January and February. Late winter and spring

feeding on natives was scattered. In April and May of both years squirrels fed on old pine

cones, most often digging them up, but occasionally removing them from the tree. Cypress





89

remained a minor element in the diet in all winter and spring months of 1996 and 1997,

except February 1996.

Fox squirrels fed on large hypogeous fungi from February through October. While

they fed on both the mycelia and the fruiting bodies, the most readily observed behavior

was the feeding on the large fruiting bodies. The peak season for observation of fungi

feeding varied slightly, June and July in 1996 and May and June 1997. Fungi feeding was

concentrated in patches that had deep litter layers, generally beneath pine and cabbage

palm, but also in stands of cypress and pine. Of the 111 fungi feeding observations at 53

tracking points, 86% were in litter, 9% in grass, and 5% in trees. A squirrel might carry a

fruiting body up a tree, where he would hang upside down and eat. In peak season, I

recorded periods of concentrated feeding when a squirrel would eat the large caps (5-8

cm) of 4-5 fruiting bodies within 10 minutes.

In May and June 1997 squirrels fed on concentrated patches of Mahogany

webworm larva, Macalla thyrsialis (John Heppner, personal communication), buried in

soil, grass or litter. They bit the tip from the cases and pulled the small caterpillars out

with their teeth. During the same period they showed no interest in the tent caterpillars

that rained to the ground from the infested cypress trees.

In both 1996 and 1997, March through May were periods of concentrated feeding

on non-natives. In March 1996 to June of that year, over 40% of recorded feeding was on

bischofia (Bischofiajavanica), an Asian species that forms clusters of small, dark berries.

Squirrels initially fed in trees and then moved to ground feeding, where they gathered in

groups of 2-8 to feed on the fallen fruit. Four large bischofia trees on the Cypress course






90


drew squirrels in mid morning and late afternoon feeding groups. In 1997, levels of

bischofia feeding were lower, with peaks in February through April in the 20-30% range.

Bottlebrush trees, native to Australia, drew squirrels to feed on their spikes of

scarlet flowers in March and April of both years. Silk oaks (Grevillea robusta), another

Australian native, with their orange-fringed flowers, also attracted fox squirrels for flower

feeding. In 1996 the peak feeding was late April and May, in 1997, late March and April.

Queen palms, common throughout Royal Poinciana, produce a bright orange,

aromatic, 2 cm oval fruit. A squirrel feeding on the fruit makes a distinctive, loud grating

sound, whether hanging from clusters of new fruits or digging up a previously buried

specimen, and so identification is easy. Queen palm feeding was high in February of both

years and moderate in April and July of 1996 and May of 1997. Squirrels regularly buried

queen palm fruits and most of the spring 1997 records are squirrels feeding on fruit they

dug up. Heavy trimming of palms in the fall and winter of 1996-97, removed most of the

available fresh crop. Regrowth and new fruits did not appear until summer of 1997.

Ten large ficus trees (Ficus spp.) dotted the Cypress course and stands of large

ficus were common at Hole in the Wall. These drew squirrels for feeding from January

through August. A variety of species with staggered fruiting times produced the most

concentrated feeding in May through July.

Fruit of the tallow tree (Sapium sebiferum) and Java plum (Syzigium cumini) trees

were feeding sites for short seasons. Tallow trees provided feeding in January and

February and Java plum in June and July. Feeding on Java plum was high in July 1997 as

the study ended, when squirrels gathered in 4 stands that produced aromatic crops of fruit

on the Cypress course.





91

The 1996 fall feeding on native species was the period with the lowest food

diversity. In both years, diversity was higher in February through July. Diversity was

noticeably higher in 1997, February to April and June, when a broader range of non-native

species and a greater use of native species were seen in the feeding records. This

difference was not only a result of changes in who was collared, but was reflected in the

expanded diets of individuals collared both seasons. While 4 squirrels showed

concentrated bischofia feeding in spring 1996 (8-23 observations), none of these repeated

the pattern in 1997. All increased the number of species they used. Only 1 individual,

subadult ROPO 27, showed concentrated bischofia feeding in spring 1997 and she

regularly fed on the 2 large bischofia trees near her birth tree. Her mother, ROPO 2, fed

on at least 10 species each spring.

The feeding data at Site 2, covering January to July 1997, were aggregated due to

low numbers (Table 12). Feeding on non-natives was dominant, as it was in the late winter

and spring at Site 1. Native foods included pines, larvae and mushrooms. Pine feeding,

16% of total, was primarily cones from previous seasons. Larvae of the same species as

Site 1 and mushrooms accounted for an additional 16% of feeding records. Cypress were

a minor element of the diet and not common at Site 2 or neighboring courses. Sixty-seven

percent of the recorded feeding was on non-native sources, while 33% of the sightings

were squirrels feeding at feeders. Ten feeders were available to fox squirrels, whether

placed there for their use or for birds. Eight of the feeders were on Royal Palm property or

adjoining residential property, 2 were on the west side of the neighboring Hibiscus course

(Figure 25). Bottlebrush trees and queen palms provided feeding opportunities at Royal




Full Text

PAGE 1

POPULATION ESTIMATES, HABITAT REQUIREMENTS, AND LANDSCAPE DESIGN AND MANAGEMENT FOR URBAN POPULATIONS OF THE ENDEMIC BIG CYPRESS FOX SQUIRREL (Sciurus niger avicennia) By REBECCA SELFRIDGE DITGEN A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 1999

PAGE 2

Copyright 1999 by Rebecca Selfridge Ditgen

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Dedicated to my father, who first took me outdoors and shared his love for wild things, and in loving memory of my mother, whose life of gentle strength and patience continues to inspire me.

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ACKNOWLEDGMENTS I thank the Florida Fish and Wildlife Conservation Commission, Nongame Wildlife Division, for their continuing attention to the pressures and consequences of rapid landscape change in southwest Florida. I am grateful for their funding and the support which made this project possible. I thank the staff of the Florida Museum of Natural History for administration of the funding. I thank my committee for their time and valuable insights. Dr. Stephen Humphrey has given greatly appreciated support and encouragement, and with patience and generosity has been the advisor I wanted and needed. Dr. John Eisenberg has shared his rich knowledge of mammals and, as teacher and friend, has inspired me to reach for the best in myself. Dr. Mel Sunquist has enthusiastically shared his skills in fieldwork, his hallmark sense of humor, and the joys of field classes at the Ordway. Dr. George Tanner has provided an excellent balance between attention to detail and the long view, as well as the important sense of perspective we often forget in the midst of all of this. And Dr. Mark Brown has been gracious in his willingness to put up with an idealistic wildlife biologist. My work relied on access to the private property of over 60 golf courses in Lee and Collier counties. I thank the golf course superintendents, staffs and members of these courses for their generous contributions to my work. Special thanks and deep appreciation goes to the 2 clubs that facilitated the radio-telemetry studies centered on their property. At Royal Poinciana Golf Club, Gary Grigg, CGCS, and his staff provided assistance and iv

PAGE 5

access for 2 years of work, while Dale Walters, CGCS, and his staff at Royal Palm County Club assisted in my work for the final year of the field study. Courses adjoining these main sites also permitted frequent visits to locate collared fox squirrels. These clubs included Hole in the Wall, Wilderness, Country Club of Naples, Quail Run, Bear's Paw, Hibiscus, and Royal Wood. I would also like to thank Tim Hiers and Mike Mongoven for their contributions to my understanding of the management and history of golf courses in southwest Florida. Hardy thanks go to colleagues in the field and office. Joys and challenges of trapping and collaring were shared by 2 excellent field assistants, Audrey Grieser and John Pamilio. Jay Harrison helped with the analysis of the data on the 60 course sites. John Shepherd gave generous assistance with vegetation sampling and mapping, and with GIS quagmires. For lively discussions, timely and invigorating e-mail, and a shared passion for work in the mammal world, thanks go to Lisa Molloy, Susan Walker, Deb Jansen, Kae Kawanishi, and Candace McCaffery. I thank the 38 fox squirrels whose fondness for peanut butter and pecans led them to carry radio-transmitters for up to 20 months as they moved, mated, fought and played. For support, understanding, and the courage to marry a woman in the final months of writing her dissertation, I thank my loving and wonderful husband, John. v

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TABLE OF CONTENTS ACKNOWLEDGMENTS iv ABSTRACT viii INTRODUCTION 1 Framework 1 Problem 3 STUDY AREA 7 Landscape Study 9 Radio-Telemetry Study 9 METHODS 17 Landscape Evaluation and Censusing 17 Sampling 17 Data Analysis 19 Habitat Use and Demography 20 Radio-telemetry 20 Vegetation Sampling 24 Data Analysis 26 RESULTS 30 Landscape Evaluation and Censusing 30 Squirrel Counts 30 Course Attributes 30 Landscape Evaluation Index 34 Demographics and Habitat Use 37 Survival 37 Reproduction 41 Mortality and Disease 42 Population Estimates 43 Home Range 43 vi

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Nest Sites /z Landscape Composition and Vegetation Mapping 73 Habitat Use 82 Feeding Patterns 88 DISCUSSION 94 Higher Quality Golf Course Landscape 94 Lower Quality Golf Course Landscape 1°1 Landscape Evaluation l^ 4 Summary MANAGEMENT RECOMMENDATIONS 1 12 APPENDLX A, TREE SPECIES ON STUDY SITES 117 APPENDIX B, GOLF COURSES, SQUIRREL LEVELS, AND LEI RANKINGS .... 1 19 APPENDIX C, HOME RANGE DATA FOR SITES 1 AND 2 121 LIST OF REFERENCES 125 BIOGRAPHICAL SKETCH 132 Vll

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Abstract of Dissertation Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy POPULATION ESTIMATES, HABITAT REQUIREMENTS, AND LANDSCAPE DESIGN AND MANAGEMENT FOR URBAN POPULATIONS OF THE ENDEMIC BIG CYPRESS FOX SQUIRREL {Sciurus niger avicennia) By Rebecca Selfridge Ditgen December 1999 Chairman: Dr. Stephen R. Humphrey Major Department: Wildlife Ecology and Conservation The Big Cypress fox squirrel (Sciurus niger avicennia) is endemic to open forests of southwest Florida. Rare in wild lands of southwest Florida, its remains on certain golf courses in Lee and Collier counties. This study was carried out from August 1995 to December 1997 to document squirrel population levels in a variety of golf course landscapes, to determine habitat use and requirements of course populations, and to provide guidelines for favorable landscape design and management in developing areas. Elements of vegetation composition and structure, and landscape configuration were recorded at 60 golf courses. Counts of squirrels were made at each course to determine fox squirrel population levels. A Landscape Evaluation Index, developed from viii

PAGE 9

cluster and factor analyses of landscape elements, allowed ranking of the 60 courses in terms of their suitability for fox squirrels. Radio-telemetry was used to examine home range size, habitat use, and population dynamics at 1 high quality course and 1 lower quality course. Tracking studies indicated a density of 42.4-49.8 squirrels/km 2 at Site 1 and a density of 6.3-8.2 squirrels/km 2 at Site 2. Squirrels fed heavily on pine and cypress from late summer to mid-winter, and relied on native and exotic species between March and May. At Site 2, squirrels showed a heavy reliance on feeders between January and July. The Index identified 7 courses with high quality landscapes. All were part of 36 hole courses, contained large stands of open pine and cypress, and had large contiguous areas free of automobile traffic. The remaining courses had unfavorable landscape elements: isolation within developed landscapes, small stands of undesirable species, heavy understory vegetation, and complex development patterns. Twenty-three courses offer little opportunity for habitat improvement. Thirty courses can improve habitat for present fox squirrel residents, but do not contain the landscape features required for long-term populations. Landscape design and placement are crucial in creating and preserving fox squirrel habitat. Courses, or groups of courses, must contain large areas free of roadways and development to allow safer movement within large home ranges. Vegetation must include large stands of pine, cypress, cabbage palms, and associated native trees with open understories. ix

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INTRODUCTION Framework In the past decade, questions of ecologically sound landscape design and management have attracted the attention of an increasing diversity of scholars and practitioners. Researchers in wildlife and conservation biology, landscape architects, planners, developers, and a range of physical and biological scientists have worked to integrate a growing body of biological principles and physical science with species and community ecology to manage landscapes for diversity and sustainability (Forman 1995, Saunders et al. 1991, Soule and Kohn 1989, Turner 1989). With an understanding of the rapid rates of land conversion and a cognizance of perceived human needs, the visions of design and management reach from large-scale regional preserves (Carr et al. 1994) and statewide, single species management plans (Mech 1998, Stith et al. 1996), to smaller scale, countywide urban parks (Mazzotti and Morgenstern 1997) and woodlot planning (Fitzgibbon 1993). Within this larger movement toward ecological design and management, stirrings of interest have emerged in some previously untouched arenas. One of these is in the community of golf course designers and managers, where individuals have begun to search for ways to create more ecologically responsible golfing developments (Grigg 1990, Foy 1989). Golf courses, considered by opponents of development to be the antithesis of 1

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2 ecological diversity and sustainability, have been portrayed as destroyers of wildlands and conduits for pesticides and fertilizers (Edmundson 1987, Foy 1989, Tietge 1992). In response to such criticism and because of personal and professional interest in a more diverse environment, advocates for change are attempting to move golf courses and accompanying developments toward more ecologically sound designs, to create more "naturalistic landscapes", and thus to encourage the preservation of native plants and animals (Dodson 1990, 1994, Leuzinger 1994). To date, much of the effort toward responsible golf course design and management has focused on badly needed reviews of turf systems and course facilities, as well as broader looks at maintenance of native vegetation in roughs (Balough and Walker 1992, European Golf Association Ecology Unit 1995, Weston 1990, 1994). Scientific research examining the benefits to wildlife of more natural and diverse golf course landscapes is just emerging. Initial research addresses the role these newer or more "naturalistic" courses may play as habitat for birds. Early work indicates they may contain more species than surrounding landscapes of agriculture or dense development (Terman 1997). Because even the most sensitively designed golf courses will save native vegetation in patches separated by manicured, exotic grass, and generally contain private homes and accompanying vehicle traffic, they cannot be expected to favor bird species easily disturbed by humans or those in need of large areas without edges (Moul and Elliott 1994, Terman 1997). Recent work in Kansas has shown that more natural courses preserving large remnants of native vegetation may create an avian habitat intermediate to dense development and wildlands (Terman 1997).

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3 While there is a growing interest in understanding and improving golf courses as habitat for avian species, little research has been undertaken to examine their ability to support mammals or other non-avian wildlife, nor has research considered the impact of surrounding landscapes on golf course wildlife (Terman 1997). Work by Jodice and Humphrey (1992) in southwest Florida suggested that the threatened Big Cypress fox squirrels may occur at higher densities on golf courses near the coast than in preserve lands to the east (Jodice and Humphrey 1992, 1993, Maehr 1993). An investigation of golf courses as potential habitat for these relatively small, though wide-ranging mammals, offers an excellent opportunity to address the broader question of golf course landscapes as habitat for non-avian wildlife. Problem Fox squirrels (Sciurus niger) are a diurnal, arboreal species inhabiting open forests of the eastern and central United States (Hall 1981). The 4 subspecies of the southeastern states are larger and more varied in color than those to the north and west and prefer open pine forests with oaks and associated hardwoods (Kantola and Humphrey 1990, Moore 1957, Weigl et al. 1989). Of these, the Big Cypress fox squirrel (S. n. avicennia) is the most restricted in geographic range, found only in the southwest tip of Florida, south of the Caloosahatchee River and west of the true Everglades. Native to open stands of slash pine (Pinus elliottii), cypress {Taxodium spp ), and tropical hardwoods, these squirrels frequently feed and move on the ground. Their relatively large size and habits of ground use make them especially vulnerable to the widespread landscape changes promoted in recent decades (Humphrey and Jodice 1992, Moore 1956, Williams and Humphrey 1979).

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4 Human activities affecting fox squirrel populations are widespread and varied in southwest Florida. Changes in fires cycles on large preserves and privately owned forests have allowed development of heavy understory vegetation not conducive to fox squirrel movement and ground feeding. Conversion of range lands to citrus groves in northern and central agricultural areas has eliminated open, parklike habitat favorable to fox squirrels (Pearlstine et al. 1997). Rapid urbanization of coastal property in Lee and Collier counties, from Naples to Ft. Myers, has created fragmented habitat with serious obstructions to squirrel movement, resulting in isolated populations amid shrinking green space (Moore 1954, Williams and Humphrey 1979, Jodice and Humphrey 1993). Demographic trends in Lee and Collier counties, two of the fastest growing counties in Florida, clearly illustrate the forces driving land conversion in the coastal zones of both counties. The 1970 permanent population of Collier County was 38,040. Between 1980 and 1990, Collier County grew by 77% (Collier County DESD 1996). In 1995, Collier had a permanent population of 197,400 and a seasonal population of 245,000. Projections indicate it will grow to between 508,00 and 770,00 by 2020 (Collier County 1996). The 1999 population of Lee County is given as 410,000 and is expected to reach 940,800 in 2020 (Lee County DCD 1998). Most of the development in both counties will be concentrated along the coast, with Collier County expecting full development west of highway 951 by 2050 (D. Weeks, person, commun). While fox squirrel populations have apparently declined in preserves such as Big Cypress and Corkscrew Swamp and have vanished from dense housing developments and commercial areas, they remain on certain golf courses within and near the burgeoning developments of western Lee and Collier counties (Deborah Jansen pers. commun., Jodice

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5 1990, Jodice and Humphrey 1992, 1993). Golf courses with remnant open pine and cypress stands preserve fragments of suitable habitat within a swirl of traffic and commerce. In an effort to understand the ecology of these golf course fox squirrel populations, Jodice (Jodice and Humphrey 1992) undertook a study of their diet and activity patterns on 4 Naples courses in 1989-1990. His work relied on focal-animal sampling of individuals located visually rather than by radio-telemetry, undoubtedly allowing bias in a species that is often difficult to see. His work successfully highlighted questions of population levels and suitability of golf course habitats (Jodice and Humphrey 1993, Maehr 1993). It became clear that little is known about spatial needs, movements, habitat requirements, or demography of these urban populations and still less about the prevalence of these golf course populations and the landscapes that might promote their survival. To evaluate the usefulness of golf courses as refugia for Big Cypress fox squirrels we must know more about the ecology of the species as well as the impact of landscape features and configuration on their survival. Can they feed, move, reproduce, and survive for sustained periods within the fragmented habitats found on golf courses? Are all courses suitable or just a few? And why? This study will address these questions by looking at home range size, habitat use, feeding patterns and demography of fox squirrels on golf courses and by identifying golf course landscape features favorable to their survival. The specific goals are as follows:

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6 1. To survey the status of golf course populations and the landscape elements of a range of golf course types in order to determine which features favor their survival. Sixty courses in western Lee and Collier counties will be considered. 2. To gather data on home range size, dispersal, habitat use, and population dynamics through the use of radio-telemetry. Two golf course populations in Collier County will be studied, one with high numbers of squirrels and one with lower numbers of squirrels. 3. To evaluate the potential of the 60 golf courses as refugia for urban populations of fox squirrels. 4. To provide recommendations for design and management of golf course landscapes to improve habitat for Big Cypress fox squirrels.

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STUDY SITE All study sites were located in the western half of Collier and Lee counties in southwest Florida (Fig.l). The area has a humid subtropical climate with heavy influence from the surrounding warm waters and the seasonal changes in the Bermuda high (Chen and Gerber 1990). These features give rise to cool dry winters and warm, rainy summers and autumns, with extreme events such as occasional hard frosts and hurricanes playing a strong role in the composition of the vegetation community. Native vegetation of the flatwoods physiographic region in which the sites were located includes pine flatwoods, cypress domes, and mangroves. The presence of Entisols, Histosols, and Spodosols reflect a mixed terrain of high, relatively dry, sandy ridges, and low, poorly drained swamps (Brown et al. 1990). Patterns of temperature and precipitation varied from year to year in the 3 calendar years of the study, with wide deviations from normal in summer precipitation (NOAA, 1995-97). Summer and fall of 1995 were extremely wet. Stations at Ft. Myers and Naples reported 1 .7 m or more of precipitation between June 1 and October 31, more than 0.76 m above normal. Flooding and long-term standing water were common on most sites during late summer and fall 1995. In the same months of 1996 the stations received only 0.56 m of rain, a 0.36 m deficit. January 1997 to August 1997, when the tracking studies ended, had normal levels of precipitation. The winter of 1995-1996 had at least 2 cool periods, with 4 nights of 0 C , and widespread damage to the more tropical flora. The winter of 7

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Figure 1 . Location of study sites in western Lee and Collier counties. Black dots are clubs visited for landscape and squirrel surveys. Red dots are sites of the radio-telemetry studies.

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9 1996-1997 was warmer than normal. An average January, with warm weather (19 C) and 1 light frost, was followed by 2 months of high temperatures. February averaged 22 C, 3 .5 C above normal, and March 24 C, 3 C above normal. Landscape Study Of the 60 golf courses selected for the landscape analysis and fox squirrel censusing, 18 were in Lee County, south of the Caloosahatchee River, and 42 were in Collier County. Course landscapes ranged from undeveloped, with large tracts of native vegetation, to intensely developed courses having close-set, multiple story condominiums on both sides of the fairways. Highly developed courses usually allowed for few trees, native or exotic, in the roughs. Courses ranged in age from over 40 years to those recently opened and still under development. The oldest courses in the study, located near the Gulf Coast, commonly were isolated from other clubs and were surrounded by development. On the eastern edge of development courses tended to be newer, often grouped together, and were located within a mixture of increasing development and remnants of pine and cypress stands. Radio-Telemetry Study Two 18-hole golf courses in Collier County, Florida, were selected for the radiotelemetry studies (Figs. 2,3). Site 1, the 18-hole Royal Poinciana Cypress Course, was half of a 36-hole private Royal Poinciana Golf Club built in 1971 in central Naples. Royal Poinciana has no residential development within the 135 ha of the golf course grounds. Fairways are bordered by open stands of moderate-size pines (Pinus elliottii var. densa),

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10 Figure 2. Site 1, location map. Identification of golf courses known to be used by fox squirrels from Site 1, Royal Poinciana Golf Club, Cypress Course. Five courses adjoin Site 1, including the Pines Course in the same club. Bear's Paw is south across Golden Gate Parkway. Stippled area is pine forest.

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11 Figure 3. Site 1, aerial photograph. Configuration of Site 1 and the surrounding landscape. Royal Poinciana Cypress Course is located in the center, with 6 other 18-hole courses nearby. Names on Fig. 2. r

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12 cypress (Taxodium ascendens and T. distichum) and cabbage palms {Sabal palmetto), and plantings of non-native broad-leafed evergreens (Appendix A). Automobile traffic around the Cypress Course is limited to a short segment of private entrance roadway on the north side of the Cypress Course, with no public roadways on the boundaries of the course. Three golf courses, including the 18-hole Pines Course within the same club, comprise the western, northern and part of the eastern boundaries. The south boundary is an undeveloped pine stand of 1 15 ha and the remaining eastern boundary is residential development of varying density. Royal Poinciana and the neighboring clubs are located within a tract of approximately 1020 ha containing 6, 18-hole golf courses, of which 3 are undeveloped, 2 moderately developed, and 1 heavily developed. The tract contains 230 ha of forested land, ranging from drier pine to swampy cypress stands. The 1020 ha tract is bordered by 4 extremely busy roadways, Goodlette-Frank, Pine Ridge, Airport-Pulling, and Golden Gate Parkway. Known predators at the site included eagles, bobcats, great horned owls, raccoons, rat snakes, and the club house cat, which was allowed to roam the course at night. Site 2, Royal Palm Country Club, is a developed 18-hole course near the eastern limit of intense suburban development along Highway 41 East (Figs. 4 & 5). The club, built in 1970, and the adjoining housing development cover 150 ha, of which 75 ha are private homes, condominium property and roadways. The club lies within a landscape currently undergoing rapid and dense development. All the fairways have development on both sides, either single family homes or large condominiums. When I began research at this site in 1996, 8 undeveloped, pine-covered lots remained along the fairways. In the next 12 months at least 4 of these were developed, with all of the pines being cut. The

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13

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15 dominant vegetation of the Royal Palm roughs and the surrounding condominiums is open pine stands. Non-native trees of a limited variety are scattered around the course and are common in the small lots of the surrounding private homes. Cypress trees are present but are not common and generally are small. The southwest boundary of Royal Palm adjoins the public 18-hole Hibiscus Golf Course. The Hibiscus course and surrounding dense development, together 132 ha, have few native tree species and narrow, open roughs with only small and scattered stands of trees (Appendix A). Busy 2-lane and 4lane roadways border Royal Palm on the north, east and south, and Augusta Boulevard bisects the course from north to south. Known predators at the site included eagles, raccoons, and a domestic cat. Activity on the courses changed with the seasons. Golf play was heavy on both courses from December until April, with Royal Palm frequently having play on every hole from 0730 until late afternoon. Royal Poinciana was generally less crowded. Summer play was light, with each course closing 1 day a week for intensive maintenance work on the course and roughs. Daily maintenance work began at 0530 or 0600 and continued until late afternoon. In winter, maintenance crews worked on the courses every day, 0630 until 1430 weekdays, and 0630 until noon on weekends. Maintenance for the removal of vegetation in the understory of tree stands included mowing, hand removal of shrubs and herbaceous plants, the addition of pine straw and the use of herbicides. Both clubs irrigated the fairways, greens and tees in the early morning and often again in the early evening. Cone production on the study courses was noticeably higher in 1996 than 1997. In the summer and fall of 1996, following a wet 1995 summer and fall, cone production on

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16 both pine and cypress trees appeared to be high, with both species heavily laden throughout the Cypress Course. In 1997, little cone production was evident on pines and the cypress suffered an infestation of tent caterpillars in the late spring, resulting in widespread defoliation of cypress and lower early season cone production.

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METHODS Landscape Evaluation and Censusing Surveys of golf course landscapes and fox squirrel populations were conducted to determine the status of fox squirrels and to identify landscape features favorable to their survival. Sampling Sixty golf courses in western Lee and Collier counties were selected for landscape analysis and squirrel censusing. In selecting the courses I looked for a range of landscape types relating to • density of development surrounding the club boundaries, from heavily developed to undeveloped; • type and configuration of development on the course, from undeveloped, nonresidential courses to those with dense development of houses and/or condominiums; • character of the rough vegetation, from little and scattered, through open tree stands, to dense forest stands with heavy understory. To obtain pre-visit information about age and landscape configuration for many of the 80+ courses in Lee and Collier counties I interviewed Tim Hires, Collier's Reserve Country Club, Naples, and Mike Mongoven, Ft. Myers and Eastwood Golf Clubs. I selected 66 courses for examination and contacted the courses through their superintendents of maintenance, individuals frequently most familiar with the landscape 17

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18 activities and wildlife on the courses. Superintendents proved to be excellent, and often interested, sources of information and influence. I made introductory visits to a majority of the 66 courses in August 1995December 1995. Exceptions were made for courses flooded by heavy summer and autumn rains. Three clubs did not wish to take part in the study; from the remaining 63 courses I selected 60 courses in 47 clubs, 42 in Collier County and 18 in Lee County (Appendix B). Each club in the study was visited 3 times for censusing fox squirrels and landscape analysis. The exceptions were those courses at which I did not see fox squirrels or sign of fox squirrels on the first visit and the superintendent and course workers had not seen fox squirrels for at least 1 year. At these courses I made no more than 2 visits for landscape analysis and squirrels censusing. Squirrel censusing took place between September 15 and May 15 each year, as squirrels were less active on the ground in the summer. Squirrel counts were conducted mornings, 0700-1030, or late afternoons, 15001730, when squirrels were most likely to be on the ground and visible (Jodice and Humphrey 1993). Sampling times were limited to sunny days with light or no wind and temperatures over 18° C. I sampled for squirrels by driving around the course in reverse order, to decrease my interference with golfers. I stopped at each fairway for 10 minutes and selected 1 or 2 locations most likely to attract squirrels: open tree stands, trees with food items, and feeder areas. I searched trees and areas with open ground while listening for sounds of movement and communication. I recorded squirrel sign, nests, cone middens, and palm leaf or bark pealing. I recorded each squirrel spotting and did not attempt to determine if animals were recounted after moving to another area of the course. I counted individuals

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19 on private property adjoining the fairways, such as those using feeders. The highest count of the repeated visits was recorded as the number of squirrels seen at that course. Five categories of squirrel counts were created to allow grouping of courses by population levels. They were • Level l~none present, a course at which I did not see fox squirrels and the superintendent and workers had not seen them for over a year; • Level 2~none seen, a course at which I did not see fox squirrels but the superintendent reported sightings within the past year. This often meant traveling squirrels were on the course for a few weeks or 1 or 2 come from neighboring courses for occasional use. • Level 3 —low, the highest number of squirrels seen on the course was 1-5. • Level 4~medium, the highest number of squirrels seen on the course was 6-10. • Level 5— high, the highest number of squirrels seen on the course was 1 1 or more. In the evaluation of habitat variation between courses I was interested in landscape features that could impact squirrel feeding, movement, nesting or predation. With these needs in mind I collected data on course configuration, place in a larger landscape, vegetation composition and structure, predators, course history and human interactions with squirrels. A field survey of questions that could be answered with a yes or no response was developed to report on this range of landscape attributes. All questions could be completed during a 3 hour tour of an 18 hole course and a 20 minute interview with a knowledgeable superintendent. All final landscape surveys were conducted between April and December 1997. Data Analysis Fifty-one responses, or attributes, from the landscape surveys were used to examine landscape variation among the 60 sampled courses. Two methods of examining variation were used. Single linkage cluster analysis was used to identify aggregations of

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20 similar courses based on 51 landscape attributes (Statistical Analysis System 6.0). Two prominent clusters were selected for further examination. Chi-square (a=0.05) was used to test the hypothesis that attributes were randomly distributed between these clusters. Factor analysis was used as another method to identify factors or groups of attributes that explained variation among the courses (SPSS). The 29 distinguishing attributes identified though cluster and factor analyses were used to create a Landscape Evaluation Index (LEI) (Bender et al. 1996, Brooks 1997, Reading et al. 1996, Thomasma et al. 1991, USFWS 1980, 1981). The Index allowed ranking of all courses and a comparison to squirrel population levels. These attributes were grouped into 3 components according to the landscape feature they described: 1) vegetation, 2) ground cover, and 3) landscape position. Attributes were weighted according to the differences in their frequency between high and low cluster courses and their ranking in factor analysis. For each course, the sum of attribute weights was expressed as a fraction of the sum of weights for an ideal course. A score of 1 indicated that a course had all of the desirable characteristics; a score of 0 indicated that it had none. The geometric mean of the 3 component scores was taken as an overall LEI (Reading et al. 1996). Courses were then ranked according to the Index and compared to patterns of squirrel sightings, as indicated by the 5 levels of population (Van Home 1983). Habitat Use and Demography Radio-telemetry A radio-telemetry study was conducted to gather data on home range, dispersal patterns, habitat use, feeding patterns and demography of two fox squirrel populations.

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21 Two sites were selected for the study, one without residential development, containing a high number of squirrels, and another, well developed, with lower numbers of squirrels. Criteria for selection required that the sites have: 18-hole courses with similar size fairways and roughs, prominence of open pine stands, a course configuration that allowed movement around the course throughout the day, absence of 3 highly invasive exotic tree species (Casuarina spp., Schinus terebinthifolius, and Melaleuca quinquenervia), squirrels that were not fed by golfers and therefore not tame, and the strong support of the course superintendent and the club greens committee for the research project. The last was especially important as the clubs would provide golf carts for 12-20 months of tracking endeavors and would support the regular presence of a non-member researcher and collared fox squirrels on their courses. Permission to work on Royal Poinciana Cypress Course was granted in November 1995 and that for Royal Palm in July 1996. Trapping took place during 4 periods: Site 1 only, December 1995-March 1996; Sites 1 and 2, JulyAugust 1996, November 1996-February 1997, and July 1997. Because of the public nature of the trap sites and the desire to decrease stress to individuals, especially females who might be pregnant or nursing, a trap line was not used. Instead, squirrels were trapped using a focused trapping method in which 1 or more traps were set for 1 or 2 specific individuals in a small area. One-ended Tomahawk #204 squirrel traps were baited with an oily, natural peanut butter and pecans. Traps were placed on the ground under trees where squirrels were feeding or resting or within 7 meters of individuals feeding on the ground. Often the traps were covered with Spanish moss or palm leaves. The traps were baited, set and covered with moss at some distance from the trap site and rapidly dropped off from a golf cart. Squirrels were acclimated to carts and

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22 would return to the place of feeding after I moved away from the trap. I watched the traps from 30-50 m. In the season of low food supply, squirrels could frequently be baited into the traps within 10 minutes to 1 hour. This method was generally successful during the winter months, when 2 or 3 individuals might be trapped and collared in a day. In the summer and autumn months, particularly in food-rich 1996, squirrels were extremely difficult to trap. To ensure recapture and collar removal at the end of the study in July 1997, individuals were baited with oily peanut butter and pecans for 2 weeks prior to the trapping period as they were located during normal radio-tracking. Final trapping was further aided by an apparently lower pine and cypress productivity in 1997. Trapped squirrels were covered and moved to the cart within 1 minute. Removal of the trap cover encouraged them to move into a dark cloth and net restriction tube that was attached to the opening end of the trap. While constrained in the bag they were weighed and given an injection of Ketamine HC1 (lOOmg/ml) in the hip. Individuals 675800 gms were given 0.25cc, those 800-1000 gms received 0.3cc. After 4 minutes or when they showed little sign of movement, they were removed from the bag and were tagged in both ears with monel sequentially number tags (size #3, National Band and Tag, Newport, KY), measured, aged, fitted with radio-transmitters and photographed. Females with darkened nipples of any size were considered adult. Males with developed testes descended into the scrotum were considered adult. Males with no obvious scrotum development or with slight development were considered subadult. There was a clear difference in the pelage and scrotal development between subadult males who had never developed sexually and adult males undergoing seasonal fluctuations of testicle

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23 development. Subadults had shorter, fine fur and no vestige of scrotal development. No animals under 5 months of age were captured. Squirrels were released at the site of capture after spending 3-4 hours in a 60 x 30 x 30 cm ventilated wooden wake-up box. Rapid retrieval, covering of the trap and immediate anesthetization appeared to reduce trauma, no squirrels died during trapping or collaring procedures. From December 1995 to mid 1996, 25 gm AVM (AVM Instrument Co., Livermore, CA) radio transmitters configured as resin pods with machine belting neck bands and 6 inch back antennae were used. This model proved unsatisfactory due to repeated transmitter failure, poor service and removal by squirrels cutting the belting. In late 1996, 1 began using ATS (Advanced Telemetry Systems, Inc., Isanti, MN) transmitters with resin pods, very fine stainless steel chain neck bands and back antennae, total weight of 28 gms. These worked extremely well, with no radio failure or removal by the squirrels. Final recapture did show that 2 individuals had slight neck abrasions. Collared squirrels were located a minimum of 2 times a week, except when weather, golf course conditions or course use would not allow (Mech 1983). Individuals were located once a week in 2 of 3 daily tracking periods, 0630 to 1030, 103 1 to 1430, and 1431 to 1900 EST. Squirrels were frequently located more than once in a sampling period; data were collected on each siting. Open vegetation and ready access to trees allowed visual sighting following radio location. When a squirrel was in a nest or concealed by heavy vegetation and visual sighting was not possible, I was able to identify the tree and the area of the tree in which the animal was located. When an animal moved to another golf course I used triangulation (White and Garrott 1990) to determine its

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24 location and then I traveled to the course for visual sighting and collection of activity data if possible. When a squirrel disappeared from the course and could not be located at a neighboring course, I searched the surrounding area in all directions. In December 1996, 1 conducted an aerial search of Collier County west of highway 951 in an attempt to locate squirrels that had disappeared. Once an individual was sighted, its location was mapped on aerial photographs. Recorded data included: time, activity (3 points at 60 sec. intervals), nature of the site and location at the site, food type if feeding on identifiable material, reproductive condition if visible, number of squirrels present within 5 m (both fox and gray), and number of collared squirrels within 5 m. Records of temperature, dew point, sky condition, and wind were recorded at the start of each session. At least once a month throughout the course of the study I took visual counts of fox squirrels at each study site. I followed the procedure outlined for the 60 course squirrel counts and in addition recorded if each sighted squirrel was collared or uncollared. These counts were used to estimate the fox squirrel population of the 2 study sites. Vegetation Sampling The tree stands on three courses, Royal Poinciana Cypress and Pines and Royal Palm, were sampled and mapped to allow comparison of habitats used by the two radio collared populations. The Pines course was included in the vegetation sampling because it bordered Site 1 on 2 sides and collared male squirrels frequently used the area. The large forested stands at the Poinciana courses were sampled using a structured pattern of 20 meter diameter circular plots placed at intervals of 25 meter from center to center on a north-south line and at intervals of 30 meters from center to center in an eastwest line.

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25 This arrangement was designed for the most complete yet rapid sampling of the generally north-south tending forested areas. In the front 9 of the Pines course the pattern was oriented in an east west direction as the fairways in that section ran at right angles to the rest of the club. Within each plot, all trees over 10 cm dbh were identified and measured. All palms were counted. Presence of all saplings were recorded. Understory coverage was recorded as percentage and type and the ground cover was recorded as litter if over half of the plot had a significant layer of pine or pine and cypress litter. If not noted, ground cover was dominated by grass with occasional patches of bare soil. In the narrow or small plots of Royal Poinciana and for the entire Royal Palm course, all the trees in each discrete plot were counted. If 10 or fewer of 1 species were counted, each tree was measured. If more than 10 of 1 species was found in a stand, 10 of the trees were measured. Saplings and ground cover were recorded as in the circular plots. The Royal Palm site contained a large region of private lands belonging to condominium complexes and private homes. All pines and cabbage palms on the condo lands were counted and all trees on private lands on the 3 streets of private housing were counted. Seven courses known to be used by the squirrels of the 2 study sites were evaluated using the method designed for the 60 course landscape evaluation portion of the study. This provided data on tree species present, identification of dominant species, types of ground cover, proportion of the course in tree stands, density of the understory and types of ground cover. The large pine stand south of the Cypress course and the developing landscape east of the Pines course were not accessible for sampling.

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26 Data Analysis In January 1996, 1 conducted 10 repeat counts of fox squirrels on the Cypress course to estimate the reliability of my squirrel census technique. The counts followed the standard format presented in the landscape analysis methods section. Results of the 10 counts ranged from 8-16 with a mean of 12.4 and standard error of 0.79. Calculation of 95% confidence limits for a small sample gave a range of 10.6-14.2 squirrels (Fowler and Cohen 1992). Such a range was considered reliable for the populations with higher numbers of squirrels. Reliability in smaller populations was expected to be lower. Survival and birth rates of collared fox squirrels were calculated on 6 month intervals, as opposed to 12 months, to allow for movement of subadults out of the sample population on the Cypress course or into the adult cohort of the sample population. Only squirrels persistent in the sample population were considered survivors. Squirrels no longer persistent in the sample population included subadults who dispersed to other locations, individuals who disappeared, adults, generally males, who no longer used the study site course but were known to remain on neighboring courses, and individuals known to be dead. Survival rates for adult females, adult males, subadult males and subadult females were calculated as the proportion of the collared squirrels remaining active in the study site at the end of a 6-month period. The birth rate was taken as the number of young known to leave the nest. The survival rate for juveniles was the proportion of the summer 1996 cohort of young known to be alive at the end of 6 months, 7 of whom were part of the collared population in 1997. Most subadults, collared at around 6 months of age, moved into the adult cohort at about 12 months of age.

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27 Estimates of population size, the number of fox squirrels using the Cypress course, were taken from 13 counts of collared and uncollared fox squirrels on the course taken in the spring and summer of 1996 and spring of 1997. In late summer and fall 1996, counts were not taken because the number of collared squirrels had declined due to collar removal and squirrel sightings were extremely low during these months of concentrated feeding in pines. Procedure for the counts followed that presented in the initial description of fox squirrel censusing. From these counts I calculated the minimum know alive (MKA), the Lincoln Index, and the Bailey unbiased population estimator for small sample sizes (Bailey 1952, Krebs 1999.). I used the RAMAS Ecolab software (version 2.0, Applied Biomathematics) to estimate growth rates in the Cypress population. Estimates of the 6-month survival and birth rates of life stages were used to construct the stage-based population projection matrix. I used birth rates from the lowest and highest 6 month periods and an average of the 3 periods to generate a range of growth rates seen during the study. Tracking locations were digitized using Atlas GIS software (Strategic Mapping, Inc., Santa Clara, CA) on an overlay of air photographs (TRW-REDI Property Data, 1996) registered to 7.5 minute topographic quadrangles. Accompanying data for each point were coded and attached to point locations using Atlas GIS software. Home range was determined using the kernel method and CALHOME software (Kie et al. 1996, White and Garrott 1990, Worton 1989, 1995). Only 1 point per sampling period was used in home range analysis (Cresswell and Smith 1992). Points were selected randomly in cases where 2 or more points were recorded during 1 sampling period. The 95% contour was used to define the home range boundary of each individual and the 50%

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28 contour as the core area (Kenward 1992, Wray et al. 1992). I calculated separate home range data for year 1 (December 199 5 -October 31, 1996) and year 2 (November 1, 1996July 30, 1997) to allow for changes in the make up of the collared population. I used a two-tailed t-test for paired samples (a=0.05) to compare home range sizes of Site 1 females that appeared in both year 1 and year 2. 1 compared home range size of adult females to adult males in both years and 2 nd year subadults from Sites 1 and 2 using a Mann-Whitney test (a=0.05) (Fowler and Cohen 1990). The small number of adult females and adult males at Site 2 precluded statistical comparisons using those individuals. Habitat maps were created from aerial photographs using Atlas GIS software. All vegetation sampling plots, including condominium areas, water features, residential areas and streets, and course fairways and non-forest roughs were outlined and the area of each type was measured. I calculated basal area, density, relative basal area, and relative density of each tree species by plot. Nine categories were defined to represent the diversity of vegetation seen on all 3 courses. Vegetation categories were defined by the relative basal area (percent of total dm 2 /ha.) of pine, cypress, cabbage palm, and other native and exotic species. Tree stands were then categorized by density (stems/ha ). Plots with a pine needle litter layer or a shrub layer were identified and mapped as such. Using Atlas GIS to analyze use of the vegetation types, tracking points were overlaid on maps of vegetation classes for each course. I compared actual use of vegetation types to that predicted by the percent area of each vegetation category. Chisquare tested (a=0.05) the hypothesis that tracking points were randomly distributed among vegetation categories.

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29 Feeding patterns were examined by analyzing the tracking data in which the food item was clearly identified. All food types taken more than 5 times in the 19 months of feeding observations were considered. Changes in monthly feeding patterns were determined by calculating the percent monthly total of each species or food type consumed each month for the 19 months of feeding records. A diversity index of species use, the inverse Simpson (Krebs 1999, MacArthur 1972, Williams 1964), was used to measure both richness and evenness of use of the food types.

PAGE 39

RESULTS Landscape Evaluation and Censusing Squirrel counts I was able to sight fox squirrels at all courses that reported regular observations by course personnel. During the squirrel counts, 5 or fewer squirrels were seen at 48 (80%) of the 60 courses. Fourteen courses (23%) were level 1, with no squirrels seen during surveys and no sightings by course staff in the past year. Nine courses (15%) were level 2, with no squirrels sighted during the surveys, but course personnel reported occasional sightings in the previous year. Reported sightings on these courses were frequently traveling squirrels or an occasional visiting squirrel from a higher level neighboring course. Level 3,1-5 sightings, was the largest category with 25 courses, 42% of the total. I sighted 6 or more squirrels on only 12 courses: 6 courses (10%) were level 4, 6-10 squirrels, and 6 were level 5, with more than 10 squirrels seen. Course attributes Cluster analysis of the 60 courses with 50 attributes produced a dendrogram with a prominent cluster of 1 1 courses and a broader cluster of 18 courses (Fig. 6 ). The 1 1 course cluster, cluster 1, contained courses with a high occurrence of attributes favorable to fox squirrels and the 18 course cluster, cluster 2, contained courses with a high number of landscape attributes unfavorable to fox squirrels. 30

PAGE 40

31 c o i 6 oo 9 ? 5 in
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32 Chi square tests of the 50 attributes showed that 20 attributes were non-randomly distributed between the 2 clusters. Courses in cluster 1 were characterized by: large patches of pine and cypress trees, open understories in the tree stands, large areas of pine litter, having adjoining courses, being part of a multi-course club, and having at least 50 acres of adjoining forest. Cluster 2 courses were characterized by: a high degree of isolation, few or no sizable tree stands, low numbers of pine and cypress, no obvious dominant tree species, heavy development and busy roadways around the course (Table 1). Factor analysis produced 2 independent factors or groups of attributes explaining 27% of the variation among the courses. Factor 1 accounted for 14 .6% of the variance and factor 2 for 12.3%. Factor 1 describes variation similar to that seen in cluster analysis, large stands of open forest on larger, grouped courses vs well developed, isolated, courses with more sparse tree cover and few native species. Thirteen attributes contributing to factor 1 were the same as those showing non-random distribution between clusters in the previous analysis (Table 1). Factor 2 is a comparison of "natural" courses with a heavy understory to courses with clear understory and the presence of planted exotic trees. Eight attributes contributed to the variation explained by factor 2 (Table 1). While cluster and factor analysis were quite useful in identifying characteristics that distinguished high and low quality courses in this study, it must be noted that they do not provide a complete listing of all features that are either beneficial or harmful to squirrel survival. Attributes such as untrimmed cabbage palms, while helpful to fox squirrels were found on over half of the courses and occurred in all types of landscapes. Similarly, known predators such as domestic cats and raptors were found throughout the

PAGE 42

33 Table 1. Distinquishing attributes as selected by cluster and factor analyses. Column 1, factors which appeared non-randomly in high and low clusters. Number is the percentage difference of occurrence between high and low clusters. Column 2, an attribute in Factor 1 or Factor 2 as determined through factor analysis. Column 3, attributes are grouped as components of the Landscape Evaluation Index. Column 4, a desired answer indicates it is an attribute favorable to fox squirrels. Index weight determined by columns 1 and 2. 1 2 3 4 5
PAGE 43

34 range of course landscapes. For this reason they did not appear as distinguishing attributes of high or low clusters. The list of distinguishing characters must not be confused with a more complete listing of characters favorable or unfavorable to fox squirrels. Landscape Evaluation Index The Landscape Evaluation Index was calculated using 29 attributes, 16 in the vegetation component, 3 describing ground cover, and 10 relating to landscape position (Table 1). The 60 courses ranged from 0.0 to 0.987 on the scale of 0.0 to 1 .0 (Appendix B, Fig. 7)). Seven courses were over 0.90, 18 from 0.50 to 0.751, and 35 were below 0.50. There was a gap with no courses from 0.90 to 0.752. Examination of the landscape configuration of the courses shows that the 7 courses above 0.90 are either undeveloped or have a perimeter development plan for each course or for the entire club (Fig. 8). Courses with a LEI of 0.75 or lower generally have more complex development plans such as the shown in levels B and C in Figure 8. These require squirrels to cross streets or travel through or around larger forest stands with dense understory vegetation to move from one portion of the course to another. Index values were strongly correlated with squirrel levels (r 0.747, p«0.01). Courses that adjoin high level courses, but do not themselves have an index value over 0.90, are identified as Neighbors in Figure 7. Three of these courses have level 4 populations, though their index values would indicate a lower capacity, and 4 of the level 3 courses have higher populations than expected. Four of the level 1 courses have moderate LEI ratings, yet no squirrels. These courses are newer, developing courses at which the large forested areas have heavy, closed understory vegetation not favorable to fox squirrels.

PAGE 44

35 « i \ t • i : • i ' i t • • ;; l 1 o 5 'I "a 8 b Of) CU CU C CC 09 0) c ° 3 o 8 ON Xi £ o £ > CO cu CU * a e = 13 > cu > c o -3 3 a. cd C cu > o & 2 9 111 ° 3 5 = x « u a o '£ > w u cx S o c/a i 2 3 o > i s o CO csj CU a u u J5 cu > co x 1 ~ X! — 09 P 09 3 x 3 O o s3 O P09 In o X) X! 2P i 0) c/J J2 cu 3

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36 Al. Perimeter development A2. Concentrated interior development B 1 . Intensified perimeter development B2. Expanded interior development with preserved wetlands CI. Connected perimeters with reserved wetlands (incomplete) C2. Radiating interior development (incomplete) Figure 8. Development patterns. Configurations of golf course development, from higher quality in level A, less desirable in levels B and C. A2 is a schematic, others are examples seen in the Collier County planning maps. Each plot is 1 section, 260 hectares. Clear tan areas are the golf courses and accompanying forest stands. Green patches are reserved wetland areas. Other colors: white, to be developed residential; brown, agricultural; pink and purple, commercial. Housing areas are represented by subdivided lots and street patterns.

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37 Demographics and Habitat Use Radio-telemetry produced 2497 tracking points and accompanying records on 29 individuals at Site 1 between December 1995 and July 1997, and 254 tracking points at Site 2 between January and July 1997. These data were used to examine population structure, size and location of home ranges (Appendix C), nesting sites, habitat use, and feeding patterns. Survival Eighteen fox squirrels were collared at Site 1 in the first year, 9 males and 9 females. In the second year, 1 1 more individuals were added to the study, 4 females and 7 males. Adult male fox squirrels had lower average survival rates than adult females (Table 2). Three individuals were known dead or moved to a neighboring course as adults. Four individuals disappeared from the population and were not located again on the Cypress course or on neighboring courses. Adult females had an average 6 month survival of 0.87, the highest rate at Site 1. Two adult females disappeared in the fall of 1997. One had a home range between 2 aggressive females and was regularly chased by each. It is possible this female moved west into the Hole in the Wall and Wilderness area, though she was never seen on the courses. The second female disappeared the same day her collar was found at the base of a tree near a busy cart path. At 0.78, the survival of subadult males was higher than adult males and subadult females (Table 2). The immediate fate of all subadult males was known. In the first year of study, 3 of the 4 remained on the course and moved into the adult cohort, and the

PAGE 47

38 « ,55 c o 6 a c w o a Q JS 0-, 8 E | | g O v> 2 ^ all © »n ci ir> o \p no rOn On On On On On 8 u ' O 3 B -a J •a I o © © o o o NO NO fOn On On On On On 1(5 1 T3 CI NO O f*> f) "A 00 NO NO NO t~On On On On On On IS « 3 M NO 8 £ d o — o O O — ' NO NO t~~ On On On On On On c a Ma, a -s .a .3 . ac

PAGE 48

39 fourth dispersed to Bear's Paw course and later disappeared. In 1997, 3 of 4 subadult males survived to the end of the study and stayed on the course. The fourth died while infected with skin fungus (Dr. Sharon Taylor, personal communication). Subadult females showed the greatest change in survival rates from one season to another (Table 2). In winter/spring 1996, 3 of the 5 subadult females dispersed from the course, for a 6 month survival rate of 0.40. Two of these were found dead on other courses, 1 within the summer and the other within the year. The third dispersing female disappeared from a developed neighboring course. In winter/spring 1997, the 3 collared subadult females remained on the course in their natal home ranges though the end of the study. At that time they ranged in age from 1 1 to 13 months. None had reproduced though 1 was known to be the object of a mating chase. Trapping difficulties in the summer 1996 meant than no subadults from winter litters could be collared. All subadults collared in the winter of 1995-1996 dispersed by August, or they entered the adult cohort. The survival data for Royal Palm, Site 2, covered a shorter period of time, only 7 months (Table 3). Four adults were collared, 2 males and 2 females. All remained alive throughout the 7 months. Four subadults were collared, 2 male and 2 females. None remained on the course at the end of 6 months. One male disappeared shortly after collaring. Both females died: 1 following a severe infestation of skin fungus, the other after being hit by a vehicle on Augusta Boulevard. The surviving subadult male moved from the course through a series of progressively more distant feeding sites. At the end of the study he was living near a feeder on the edge of Hibiscus course along Highway 41 .

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40 Table 3. Survival rates for Site 2, Royal Palm, for January 1997July 1997. Survival equals persistence in the course population. No births, b(x), were recorded in the collared population between January and July 1997. SITE 2 MALESn Known dead Moved/ dispersed Disappeared Persistent in the population Survival/6 mo Adult 2 0 0 0 2 1.00 Subadult 2 0 1 1 0 0.00 FEMALES b(x) Adult 2 0 0 0 2 0 1.00 Subadult 2 2 0 0 0 0 0.00

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41 Reproduction Reproduction at Site 1 varied widely between seasons and between individuals (Table 2). Winter reproduction was lower in 1995-1996 and 1996-1997 than the summer/fall of 1996, with only 4 and 3 young known to leave the nest in the 2 cooler seasons. The 6 month rate of reproduction for the first winter season was 1.00/adult female, in the second winter season it was 0.60/adult female. Winter young were born in December through February, emerging from the nest in January through April. The summer of 1996 was one of high reproduction. Five of the 6 adult females produced evident young, with 2-4 young seen emerging from each of the 5 nests, for a total known reproductive output of 15, a rate of 2.50/adult female for the 6 months. Warm season young were born from early July to September and emerged form the nests between August and late October. Mating chases were recorded in April though July and in October. Neither female at Site 2, Royal Palm, produced young from a nest during the course of the study. One female was obviously pregnant at the time of collaring and appeared to tend a brood nest for a few weeks. She failed to show signs of long-term nursing and no young emerged from the nest. She appeared to be tending a brood nest when her collar was removed, as did the other female in the study. The surviving subadult male in the study was the offspring of one of the collared adult females, undoubtedly born in the summer of 1996. At the time of his collaring, he and another subadult regularly accompanied her. Though reproduction was not observed in collared animals during the study, some adult females were successfully reproducing at Site 2, though apparently at a lower rate than Site 1 .

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42 Mortality and Disease Known causes of death in the collared populations of Sites 1 and 2 were: automobiles (2), skin fungus infection (2), and electrocution (1). Three subadult uncollared fox squirrels at Site 1 also died from vehicle accidents, 1 from a car on the entrance road and 2 from golf carts on the course. Of the 5 squirrels known to be killed by cars or carts at the study sites, only 1 remained at the site of impact. The others moved away, sometimes several meters, before they died. Skin fungus (Dr. Sharon Taylor, person, commun), causing heavy fur loss and blackened crusting of the skin, was apparent in both populations in 1997, affecting at least 8 collared individuals. One subadult died at each site. It primarily affected subadults, though also was seen in 2 adults. The proportion of affected individuals appeared similar in collared and uncollared squirrels. One uncollared subadult died at Site 1 within a month of the collared individual. Fur loss and darkened skin as seen in a skin fungus infection was observed in squirrels at courses adjacent to the study sites prior to 1997. It was easily transmitted by contact and feeding areas appeared to be vectors, especially when squirrels fed from a concentrated food supply. At least 3 squirrels with severe fur loss and skin damage were seen sharing a feeder at CCN. The collared individuals who survived the spring 1997 skin fungus infestation regained a thick, healthy coat in the late spring molt. Information on predation is slight. A bobcat was seen killing a ground feeding, uncollared fox squirrel at Site 1 in 1997 (A Grieser, person, commun.). At least 1 female bobcat and 2 young were regularly seen in the forest areas adjacent to Poinciana on the east and west. The adult bobcat began frequenting Poinciana in daylight hours when the area immediately east of the course was being cleared for development in 1997. There is

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43 no evidence of raptor kills on the study sites, though I did find apparent raptor-killed fox squirrels at 2 other nearby courses. Population estimates Estimates of population size, derived from repeat squirrel counts at the study sites, are presented in Table 4. Range of minimum number alive (MNA) at Site 1 is 15-26. The Lincoln estimates range from 15.9-32.5 and the Bailey estimate for small samples is 15.630.5. Counts in July were understandably lower as a number of subadults had dispersed from the course and young of the season would not emerge from nests until early fall. At Site 2, the MNA range is 3-8, the Lincoln estimates range from 8.3 to 15, and the Bailey unbiased estimates are lower at 7.5 to 11.7. RAMAS Ecolab software (2.0, Applied Biomathematics) calculated an annual growth rate (k) of 0.803 using the Site 1 winter/spring 1997 reproductive rate of 0.23 young/adult for 6 months, lowest of the 3 periods. Use of the summer/fall 1996 reproductive rate of 1.25 young/adult gave an annual growth rate of 1.39. The average birth rate/adult for the 3 periods of study, 0.67, yielded an annual growth rate of 1 10. Home range At Site 1, in the first year of study, adult female home range size varied from 8.0 to 12.0 ha (x=10.1 ha) (Table 5). Home ranges of the 6 adult females cover the study site (Fig. 9) from the north end to the southern border. No other adult females were observed to make exclusive use of the course during the first year of study. A light color female occasionally fed in the northeast section of the study site, though the core area of

PAGE 53

44 Table 4. Population estimates of Sites 1 and 2. Estimates are derived from repeat censusing of each site. Tntal 1 Olal iviujvYii Collars MNA Lincoln Bailey Standard seen collars in sample Index unbiased error of Index Bailey Index SITE 1 14-Apr-96 19 16 8 11 24 27.6 26.7 4.68 28-Apr-96 7 15 3 4 18 26.3 24.0 6.00 6-Mav-96 18 14 5 13 19 19.4 19.0 2.52 25-May-96 10 13 6 4 1 Q 8 6? 6-Jun-96 11 13 5 6 18 23.8 22.3 5.09 l-Jul-96 9 13 3 6 16 19.5 18.6 3.60 l-Jul-96 11 13 2 9 15 15.9 15.6 1.92 l-Mar-97 14 19 5 9 24 29.6 28.5 4.96 20-Mar-97 8 18 2 6 20 24.0 23.1 3.86 29-Mar-97 19 18 7 12 25 28.5 27.7 4.38 6-Apr-97 15 17 7 8 24 31.9 30.2 6.32 21-Apr-97 16 17 7 9 24 30.2 28.9 5.59 ll-Jun-97 20 16 10 10 26 32.0 30.5 6.08 Means 13.6 20.9 26.2 24.9 SITE 2 ll-Jan-97 8 0 8 0 8 19-Feb-97 6 5 4 2 9 15.0 11.7 4.41 8-Apr-97 3 7 1 2 8 10.5 9.3 2.33 15-May-97 4 5 2 2 7 10.0 8.3 2.64 14-Jun-97 5 5 2 3 7 83 Z5 1.94 Means 7.8 11.0 9.2

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45 .3 3 X! < M ^ on fN O «> 00 vb sd i oo © f•— 1 pfV© Ov CN vo i— < O m h « m m >/"> On N W K 6 h t tH H rt N N H i-H f-h rs r-» On ^ On ci On On O VO ^ 1— <-H N — > i VO On. On > O Z rs I s oo |-H 00 © vci n 6 n O N 00 00 V") (*i o oo oo t O Ifi — | v© GO cn o fH h m N rON On On rs 3 8 p— < rs 00 f— CO

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46 Figure 9. Adult females, Site 1, year 1. Tracking locations and home range boundaries, defined by the 95% contour, kernel analysis, CALHOME. The summer/fall brood nest site of each female is identified by placement of her individual number.

PAGE 56

47 her home range was east of the study site in the Pines course. Of the 431 tracking locations recorded for the females between December 1995 and November 1996, only 10 were outside of the course boundaries, all but 2 of these 10 remained within the boundaries of the Poinciana. Females did not range into neighboring pine or cypress stands and only ROPO 6 crossed the entrance roadway on the club grounds to feed in a large ficus immediately north of the road. All adult females showed some home range overlap with other adult females, though none showed range overlap with more than 2 other adult females. In summer/fall 1996 all females maintained a brood nest in a location within a core area outside the home ranges of all other adult females. In the first few weeks the of summer brood nest occupancy females stayed in the nest most of the day and greatly reduced the area in which they fed (Table 6). Placement of the nest within a small mixed stand often allowed them to feed without moving to the ground. A similar pattern of reduced movement and isolation was not observed during the winter brood nesting period. Adult males at Site 1, year 1, had home ranges of 42.52 ha to 1 18.40 ha ( x = 70.84 ha) (Table 5), significantly larger than those of the adult females (MannWhitney U=0, p<0.05). The home ranges of all 4 adult males overlapped in the center of the study site (Fig 10). Other adult males were regularly seen on the course. ROPO 04 and ROPO 07 used the Cypress course almost exclusively, while ROPO 16 used all of the Cypress course and the adjoining back nine of the Pines course on the east. ROPO 17 used portions of 4 courses, the Cypress, Pines, Hole in the Wall, and the Country Club of Naples to the west. Though ROPO 17 used the open edges of large forested stands within

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48 Table 6. Brood nest home ranges. Home range size for 3 adult females occupying brood nests in summer/fall 1996, Site 1. Home range determined by kernel analysis, CALHOME. Brood nest home Brood nest core Number range, 95% contour area, 50% contour sightings ROPO 02 0.8 ha 16.7 m 2 16 ROPO03 0.3 ha 9.9 m 2 13 ROPO 06 2.1 ha 78.5 m 2 9

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49 Figure 10. Adult males, Site 1, year 1. Tracking locations and home range boundaries, defined by the 95% contour, kernel analysis, CALHOME.

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50 and between courses, he was never found in the interior of these stands. He did not need to cross the roadway to move from CCN to H1W and was regularly observed moving to HIW by crossing the canal between the Cypress course and HIW by way of a natural tree bridge on the west side of fairway 2. These tree routes were frequently used by several individuals in moving between these 6 neighboring courses separated by canals. Home ranges of the 6 collared subadults at Site 1, year 1 (Table 5, Fig. 1 1), ranged from 10.66 ha to 49.07 ha (x = 22. 16 ha). ROPO 13, a male, began making long day trips to the back nine of the Poinciana Pines course in the month prior to his dispersal. His large home range, twice as large as the mean and the next largest subadult home range, reflects these trips. The 4 subadults in the southern half of the study site show a strong overlap (Fig. 12), with all four commonly using forested areas along the 7 th and 13 th fairways. Three of these 4 dispersed to other courses between March and August 1996. The fourth stayed on the Cypress course until the end of the study. ROPO 10, a subadult female, dispersed from the southern section of Site 1 to Poinciana Pines at the end of March 1996 and in early April moved to the CCN (Fig. 13 ). Her initial movement to the front nine of the Pines course was 1 .4 km with an additional 0.85 km to CCN. She also frequented a home feeder at point B and crossed Burning Tree Drive regularly. She slipped her collar in June 1996 and was seen only once again, despite subsequent searches. ROPO 14, a subadult female, dispersed from the northeast section of Site 1 in late April 1996 (Fig. 14), moving to Bear's Paw Country Club. Her dispersal distance from her Site 1 to her first sighting at BP was 2.8 km. She occasionally fed on scraps put out at Site A with the remaining tracking sites at the edges of heavy forest on the northeast edge

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51 Figure 11. Subadults, male and female, Site 1, year 1. Home range boundaries are defined by the 95% contour, kernel analysis, CALHOME. Individuals that dispersed during the 1996 spring and summer seasons are marked (*), home ranges are those prior to dispersal.

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52 Figure 12. Four subadults, Site 1, year 1. Collared subadults in the southern half of the course. Tracking locations and home range boundaries indicate use of the course. Three of 4 squirrels, 2 females, 1 male, dispersed from the course in spring and summer 1996. Only ROPO 08 remained through year 2 of the study.

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53 Figure 13. Dispersal movement of ROPO 10. Movement from Site 1 to Country Club of Naples in late March, 1996. Movement from her central home range area to A was 1 .4 km, and from A to B an additional 0.85 km, for a total 2-week dispersal distance of 2.2 km.

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54 Figure 14. Dispersal movement of ROPO 14. Dispersal in April 1996 to Bear's Paw Country Club. The distance from her central home range area to the first off-site location, A, was 2.8 km. She was found dead at B in late July 1996.

PAGE 64

55 of the course. In July she was limping badly (L. Molden, person, commun.) and was found dead in dense undergrowth on July 24, 1996. At least 1 large cat regularly roamed the small area where she was found. In mid-May 1996, ROPO 09, a subadult female, moved from the center of Site 1 to the eastern end of the Quail Run Golf Course (Fig. 15), a heavily developed course with few tree stands, a dense street network and heavy traffic. By the end of June 1996, ROPO 09 had moved to the CCN where she often fed at a home feeder station used by 2 or 3 other fox squirrels, at least 2 of which had severe fur loss with accompanying thick, darkened skin, probably related to a skin fungus. By August, ROPO 09 was suffering from a similar fur loss when she was tracked to the center of the HTW. She returned to Site 1, Cypress course, for a brief period in September 1997. Regular contact was lost in early fall due to collar failure. She continued to use the HTW and the south end of the CCN, where she frequently crossed roadways in her daily movements. She was found dead at the side of the entrance road to CNN in April 1997. Her initial dispersal distance was 2.6 km and the total distance moved from the original home range to HTW was 6.0 km. ROPO 13, a subadult male, dispersed from the southern section of Site 1 to Bear's Paw Country Club in August 1996 (Fig. 16). On August 8, 1996, he moved from the south border of Site 1 to Golden Gate Parkway, through 1 .2 km of pine forest. The following morning he crossed the busy 4 lane roadway into BP, where he was tracked until mid August 1996. His initial 24 hour dispersal distance was 1 .9 km; his total movement distance from his original home range was 3 .5 km.

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56 Figure 15. Dispersal movement of ROPO 09, mid May 1996. First off-site location at A, a move of 2.3 km from previous home range on May 20, 1996. Located at B on May 27, 1996, at C, home feeding site on June 28, and at D on August 23. She was found dead at E, hit by vehicle, in April 1997.

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57 Figure 16. Dispersal of ROPO 13. Movement from Site 1 to Bear's Paw Country Club on August 8, 1996. Points north of A are tracking locations of 13 before that date. Movement from A to B, 1 .2 km, took 3 hours. He crossed from B to C the following morning, for a total 24-hour dispersal distance of 1 .5 km. The total distance traveled to his final sighting was 3.5 km.

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58 Figure 17. Adult females, Sitel, year 2. Tracking locations and home range boundaries, defined by the 95% contour, kernel analysis, CALHOME. Numbered site is 1997 summer brood nest of female 03 .

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59 At Site 1, year 2, the home ranges of the 5 adult females varied from 9.08 ha to 20.92 ha (x=16.40) (Fig. 17). The 4 adult females collared from the previous year, showed significant increases in home range size (Table 5) ( t=3.95, P=0.029, df=3) though each female continued to use approximately the same section of the course. ROPO 03 and ROPO 05 increased use of the edges of large forest stands on the eastern edge of HTW and ROPO 15 used open edges of forested roughs at Wilderness Country Club and a feeder area on private property adjoining the southeast corner of Site 1. ROPO 03 expanded to the north into the previous home range of ROPO 06, who disappeared in December 1996. Core areas of ROPO 03, ROPO 05, and ROPO 15 showed increases from year 1 to year 2. ROPO 1 5 and ROPO 28 had high overlap, though location of core areas indicated that ROPO 28 made heaviest use of the southeast portion of the course and ROPO 15 the southwest corner of the corner. Only ROPO 03 was tending a summer brood nest when the study ended in July 1997. The second year home range of ROPO 05, a female, had expanded to include the fall 1996 brood nest site of ROPO 03. ROPO 03 moved her 1997 summer brood nest to the north, into a tree stand formerly within the home range of ROPO 06, an adult female. The 6 adult males of Site 1, year 2, used portions of 5 courses (Figs. 18, 19). Home range size varied from 44.06 ha to 121.00 ha (jc= 90.91 ha) (Table 5), significantly larger than adult female home ranges in the second year (Mann-Whitney U=0, p<0.05). All of the males used at least some part of 2 or more courses and did so with little crossing of roadways. ROPO 20 used much of the Cypress course, visited the front and back sections of the Pines course and frequently moved into Hole in the Wall. ROPO 18 rarely visited the Cypress course, spending most of his time in the northern

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60 Figure 18. Three adult males, Site 1, year 2. Tracking locations and home range boundaries, defined by the 95% contour, kernel analysis, CALHOME.

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61 Figure 19. Three adult males, Site 1, year 2. Tracking locations and home range boundaries defined by the 95% contour, kernel analysis, CALHOME.

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62 section of the Pines course and the CCN. ROPO 07 and ROPO 08 used the Cypress course and both sections of the Pines course. ROPO 08 and ROPO 20 had the smallest home ranges, each using portions of the Cypress course and the back nine of the Pines course. Males again showed strong overlap of home ranges. Seven subadults at Site 1, year 2, had home ranges of 5.88 ha to 21 .47 ha (x=14.93) (Figs. 20 & 21). All subadults show overlap with at least 3 other subadults. None of the subadults dispersed before the end of tracking in July 1997, though ROPO 21, a male, began to use the northeast corner of Wilderness Country Club to feed in June and July 1997. Five of the subadults at Site 1, year 2, were born to adult females ROPO 02, ROPO 03, and ROPO 05 who remained on the course in the same home range areas throughout the study. Figures 22 and 23 compare the 1997 spring/summer home ranges of the collared offspring to the 1997 home ranges of their mothers. The female offspring of ROPO 02 and ROPO 03 had home ranges entirely contained within those of their mothers, while the male offspring of ROPO 03 and ROPO 05 had home ranges that extended beyond those of their mothers. The overlap of the subadult home ranges with the core area of their mother, ROPO 05, is clear (Fig. 23). Adult female ROPO 01 disappeared in December 1996 and offers no comparison with her 1997 home range area with those of her offspring, ROPO 21 and ROPO 24 . Nevertheless, it is clear that her 1996 offspring, male ROPO 21 and female ROPO 24, continued to use their natal home range, the core area of female ROPO 01 before her disappearance (Fig. 24). ROPO 24 expanded to the northeast in her mothers absence and ROPO 21 expanded to the south and west.

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63 Figure 20. Three of 7 collared subadults, Site 1, year 2. Tracking locations and home range boundaries as defined by the 95% contour, kernel analysis, CALHOME.

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64 Figure 21. Four of 7 collared subadults, Site 1, year 2. Tracking locations and home range boundaries as defined by the 95% contour, kernel analysis, CALHOME.

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65 Figure 22. Two adult females and offspring, Site 1, year 2. Females, 03 and 02, and their collared offspring from summer 1996. Thin lines are subadult offspring. Squirrels 27 and 29 are females, 26 is male.

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66 Figure 23. Home ranges of adult female and offspring, Site 1, year 2. Adult female 05 and collared male offspring 23 and 25 from summer 1996. Yellow dots indicate the core area, 50% contour, of female 05.

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67 Figure 24. Home ranges of female and 2 offspring, Site 1, year 2. Subadults 21 and 24, siblings from female 01 in the summer 1996, show overlap of home ranges and core areas. Core area of 01, who disappeared in December 1996, is shown in light blue.

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68 All 7 of the Site 1, Year 2, subadults continued to use their natal homeranges for the first year of their lives, 5 sharing with their mothers, 2 remaining after their mother disappeared. Seven of the 8 squirrels collared at Site 2 in year 2 remained to provide usable home range data. ROPA 01 and ROPA 08, adult males, had home ranges of 136. 1 ha and 303 .8 ha, respectively (Table 5). ROPA 01 used most of Site 2, Royal Palm, while ROPA 08 used Site 2 and most of the neighboring Hibiscus Country Club (Fig. 25). ROPA 08 readily moved from the east to the west end of his home range, a distance of 2.5 km, within 24 hours. Both males regularly crossed Augusta Boulevard, while their movements on either side of that busy street generally followed the fairways and appeared to minimize travel through housing. Though their home ranges overlapped, they were never seen together as males at Site 1 often were. The two adult females at Site 2 had home ranges of 13.06 ha and 30.57 ha (Table 5, Fig. 26). Adult female ROPA 04 had a home range 50% larger than any at Site 1 . She often crossed Augusta Boulevard. Adult female ROPA 06 had a home range similar in size to those at Site 1 . Her home range included a regularly stocked feeder at a private residence. On rare occasions she crossed into the central pine stand within the private housing area. Though the home ranges of these 2 adult females were widely separated, no other adult females where ever observed in the area between their 2 home ranges. Three of the 4 collared subadults at Site 2 provided data on home range size, these varied from 25.77 ha to 108.50 ha (x= 58.34 ha) (Table 5, Fig. 27), significantly larger than Site 1 subadults for the same period (Mann-Whitney U=0, p<0.05). Male ROPA 2

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69

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70

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72 was an older subadult who disappeared within a month of being collared and provides no usable home range. Female ROPA 3 used an area on both sides of Augusta Boulevard and across Forest Hills Boulevard into the Hibiscus course. Female ROPA 05 used an area on both sides of Palmetto Dunes Circle. Male ROPA 07, summer 1996 offspring of adult female ROPA 06, had an unusual home range use pattern, sequential use of small patches. He spent a few weeks to a month in a small area of a hectare or less and then moved to another area, each time moving away from his natal home range at the northwest corner of Site 2. In the final week before his collar was removed he had moved to the west side of Hibiscus Country Club along U. S. 41, 1.2 km from his natal home range. Nest Sites At Site 1 fox squirrels made regular use of untrimmed or lightly trimmed cabbage palms, bromeliads and cypress cavities for sleeping nests. Stick nests were used on occasion. The common use of untrimmed palms and bromeliads as nests eliminated the possibility of counting nests and determining the nest to squirrel ratio. Squirrels were regularly observed carrying Spanish moss {Tillandsia usneoides) to nesting sites. Squirrels rarely constructed open platform nests for daytime resting but simply draped themselves along branches. Brood nests were readily observed at Site 1 in summer 1996 as females seldom moved from the nests for 2 -3 weeks. Nests were located in mixed stands of pine, untrimmmed or lightly trimmed palms, and cypress (Fig. 9). Three females used cavities high in large cypress trees, 1 raised a litter of 4 from such a cavity, another a litter of 3. Two females raised young in the center of densely leafed palms, one a queen palm (Arecastrum romanzqffiartum) and the other a cabbage palm. The remaining female used a

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73 large bromeliad high in a large pine for her nest site. The nest was located in the base of the large plant. All females used Spanish moss in constructing and maintaining nests. The queen palm nest also contained shredded queen palm leaflets and required her to spend time stripping leaflets and tearing them into strips. Winter brood nests were similar to those in the summer of 1996 and included 2 in cypress cavities, 2 bromeliad nests, and 1 palm nest. As the study ended at Site 2, both adult females appeared to be tending brood nests. One was in a rather isolated moderately trimmed cabbage palm near a canal and the other was in a stick nest in a moderate-sized pine tree. In the winter of 1997, the same female was observed using a wood duck box with her offspring from the previous summer. All 3 nested together in the box during times of heavy rain, cooler nights, or high wind. Landscape Composition and Vegetation Mapping Landscape composition of Sites 1 and 2 and the Poinciana Pines course is presented in Table 7. While Site 2 was 2.3 times larger than Site 1, the golf course and tree stands within the roughs of the 2 courses were similar in size, though not necessarily similar in species composition or structure. The obvious difference in landscape composition between the 2 sites was the presence of housing areas at Site 2. Twenty-eight percent of Site 2 was occupied by residential development, streets and clubhouse property, and an additional 25% was condominium land and a private pine stand within a housing area. The 2 courses that comprised Royal Poinciana Golf Club, Site 1 and the adjoining Pines course, were similar to one another in general landscape composition. The smaller

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74 Table 7. Landscape composition. Landscape cover of Site 1, Site 2, and Royal Poinciana Pines course. Royal Poinciana Cypress Royal Palm Site 2 Royal Poinciana Pines Course Site 1 Area, ha %of total Area, ha %of total Area, ha %of total Total size of site 61.4 141.9 62.9 Club property in 30.0 49% 26.0 18% 24.8 39% tree stands Condominium land 0.0 0 34.9 25% 0.0 0 in trees Residential, streets 0.0 0 40.1 28% 0.0 0 ana ciuDnouse Lakes, canals, 5.5 9% 8.0 6% 5.3 8% wetlands Fairways, greens, sandtraps, driving ranges, unforested 25.9 42% 32.9 23% 32.8 52% roughs

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75 area of tree stands in the Pines course reflected the open nature of the back 9 of that course and the presence of 2 driving ranges within the course. While Site 1, Site 2, and the Pines course each had between 30.0 and 24.8 ha of tree stands or forested area, the species composition and structure of these plots were not alike. Site 1 sampling plots contained a greater diversity of tree species than Site 2 plots. (Table 8, Fig. 28,29). A mixture of native tree species, pine, cabbage palm, and cypress, dominated Site 1. Though Site 1 was high in native hardwoods, there were no class 1, pine dominant, stands. Pines were found as co-dominants with cabbage palm (11%), cypress (3%), or both (10%). Cypress was dominant in 14% of the forested area and was codominate with pines and/or cabbage palms in an additional 3 1% of the area. Palm dominant stands accounted for 8% of the area in tree stands. At Site 1, 64% of the tree stands were dominated by the pines, cypress and cabbage palms. A mixture of native species, often including oaks (Quercus virginianum, Q. laurifolia) , maple {Acer rubrum), red bay {Per sea borbonia) made up another 10% of the forested landscape. While native species dominated the landscape, the importance of exotic species on the Cypress course was seen in the extent of class 9 (mixed natives with exotics), which covered 26% of the forested area. Site 2 was dominated by pine, with 68% of the plot area in class l(pine dominant) and an additional 6% with pines as co-dominants (Table 8, Fig. 29). Cypress was a minor element of the Site 2 vegetation, dominating in only 2% of the area. Classes 6 and 7 of the mixed natives were not present at Site 2. Exotics were less common than at Site 1 , with class 9 (mixed natives with exotics) accounting for 1 1% of the area.

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76 Table 8. Vegetation classification of Site 1, Site 2, and Royal Poinciana Pines. Royal Royal Palm Royal Poinciana Site 2 Poinciana Cypress Pines Course Site 1 Vegetation classes, by relative Area, > Area, % Area, % basal area ha lid ha ha 1-Pine, 70% or more pine 0.0 0 17.8 68 3.0 12 2-Pine and cabbage palm, 3 2 11 0.9 3 2.4 10 30% or more of each 3-Palm, 60% or more cabbage 8 1.4 5 2.0 8 palm 3.5 4-Cypress, 60% or more 14 0.4 2 14 cypress 5-Pine and cypress, 30% or n o 3 0.7 3 0 0 0 more each of pine and cypress 6-Pine, cypress, c. palm, over 2.9 10 0.0 0 0.0 o 85% of the 3 combined, with each being over 20% 7-Cypress and c. palm, 30% 5.5 18 0.0 0 0.0 0 or more of each cypress and pine 8-Mixed natives-none of the 2.9 10 2.1 8 1.2 5 above with 20% or less exotics 9-M ixed natives with exotics7.8 26 2.8 11 12.7 51 not 1-7, with more than 20% exotics

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77 Figure 28. Tree stand characteristics at Site 1, Royal Poinciana Cypress. Nine vegetation classes are color coded. H= high density (>200 stems/ha), L=low density (<100stems/ha), all patches not marked H or L are moderate density (100-199 stems/ha). Presence of dense litter layer is indicated by an asterick (*); S= presence of a shrub layer. Light gray areas between tree stands are the fairways, tees, and greens.

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78 ^ * 11 ill! •g "D "B _ c o 2 2 lill 2P o ° « 3 ™ M a) u-3 a, .2

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79 Figure 30. Tree stand characteristics for Royal Poinciana Pines course. Nine classes are color coded. H=high density (>200 stems/ha), L=low density (<100 stems/ha), all patches not marked H or L are moderate density (100-199 stems/ha). Presence of a dense litter layer is indicated by an asterisk (*); S= presence of a shrub layer. Gray areas between tree sands are fairways.

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80 The Pines course had a high presence of exotic species, with 51% of the tree stand area in class 9, and a much lower area dominated by a mixture of pines, cypress and cabbage palms (Table 8, Fig. 30). Pine dominated stands (class 1) accounted for 12% of the forested area and pine/cabbage palm for an additional 10%. Cypress dominated 14% of the area. Classes 5, 6 and 7, mixtures of pine cypress and cabbage palm, were absent from the Pines course. Structure of the trees stands on the 3 courses also differed from course to course (Table 9, Figs. 28-30). Site 1 had a higher density of trees than Site 2, with 24.9 ha of forested area having more than 100 stems/ha and only 5.2 ha in lower density stands. Site 2 had a much more open landscape, with 17.9 ha of low density (< 100 stems/ha) and only 8.4 ha in high and medium levels. The Pines course was a mixed course, with a dense forested front 9 on the north and an open back 9 on the south. The presence of a litter ground cover, as opposed to grass or bare soil, and the presence of a shrub layer are noted in Figures 28-30. Site 2 had 4 plots with a significant litter layer. All were in pine dominated stands, only 1 with a high tree density. Site 1 had 8 plots with heavy litter layers, 5 of these plots had high tree density and all of these stands were dominated by a mixture of native pine, cypress or cabbage palms. Four plots at the south end of Site 1 had a shrub layer, with wax myrtle (Myrica cerifera) the most common understory species. The Pines course had 7 plots with a litter layer and 5 of these had high tree density. The more open back nine of the Pines contained 2 large plots with heavy litter layers and high to medium density of trees. The condominium areas and private forest stand at Site 2 were dominated by pines, though tree density varied widely with the number of buildings and parking lots in

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81 Table 9. Density of trees at Site 1, Site 2, and Royal Poinciana Pines. High Medium Low > 200 stems/ha 100-199 <100 stems/na stems/ha Site 1, R. P. Cypress5.2 area in ha 5.4 19.5 % -ton/ 18% /-co/ 1 / /o Site 2, Royal Palm area in ha 0.7 7.4 17.9 % 3% 28% 69% Royal Poinciana Pines area in ha 7.8 9.9 7.1 % 32% 40% 28%

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82 each plot (Fig. 29). Most condominium plots had low pine densities, with 19.8 ha in the range of 30-99 stems/ha and 5 .6 ha at < 30 stems/ha. High density pine forests (>200 stems/ha) were found in 4 plots totaling 4.5 ha and 2 plots totaling 4.0 ha had moderate densities (100-200 stems/ha). The 6 undeveloped private home vacant lots bordering the course in the summer of 1997 had dense stands of pine with scattered cabbage palms. The developed lots that circled the course on all but Augusta Boulevard had a very low density of trees, with 3 or fewer trees in most lots. Pine and queen palms (Arecastrum romanzoffianum) were the most common species. Black olive (Bucida buceras), cabbage palms, oaks and bottle brush (Callistemon rigidus) were scattered throughout the lots of private residences. Habitat Use Locations of 2138 of the 2497 tracking points of the Site 1 collared population were used to examine habitat preference on the Royal Poinciana Cypress and Pines courses. The 359 points not used were located either on neighboring courses, fairways or unforested roughs. Comparison of the aggregated area of all plots in each of the 9 vegetation classes and the number of points located within the boundaries of each class shows a non-random use of forested areas (Table 10, Fig. 31). On the Cypress course, all subsets of the population showed non-random use of the forested stands. In all cases, fox squirrels had higher than expected use of class 2 plots pine/cabbage palm co-dominated stands While class 2 plots were 1 1% of the forested area, they accounted for 18% of the tracking locations for females and 28% for males. A preference for class 2 plots was shown in both years of the study.

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83 Table 10. Habitat preference tests on tracking points at Royal Poinciana Cypress and Pines courses. Chi-square tests the hypothesis that points are randomly distributed among vegetation types. Area of each vegetation class predicts the number of points. Only tracking locations within the mapped plots of forested stands are included. Double asterisk indicates results are significant at the 0.01 level. Number of Tracking X 2 df p Points Cypress Females, both years 1206 154.08 7 5.60E-30 ** Males, both years 768 242.76 7 9.60E-49 ** Year 1, all squirrels 1036 208.12 7 2.18E-41 ** Year 2, all squirrels 938 128.51 7 1.28E-24 ** Summer 1996 276 81.57 7 6.59E-15 ** Summer 1997 340 41.29 7 7.12E-07 ** Pines All points 164 20.73 5 0.0009 ** Males, both years 140 17.59 5 0.0035 ** Year 1 all squirrels 65 7.99 5 0.1570 Year 2 all squirrels 99 18.31 5 0.0026 **

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84 ssaidX;) (juasajd l°N) 9u Id # # # # aP # in o jp o u? o «>papadxa iuojj aou3J3jjia% *r

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Site 1 fox squirrels showed consistent underuse of cabbage palm dominated plots, cypress dominated stands and plots of mixed natives. Females showed a preference for pine/cypress dominated plots, as well as pine/cypress/cabbage palm stands and cypress/cabbage palm forested areas (Fig. 31). Males underused or showed expected use of all vegetation types other than the pine/cabbage palm plots. On the Pines course the aggregation of all tracking points within the tree stands shows a preference for pine plots, mixed natives and natives with exotics. Both cabbage palm plots and cypress stands were underused (Fig. 32). Females used the Pines course so rarely, only 24 points in the 2 years of the study, that they were not examined separately for patterns of use. The summer subsets were similarly small and not separately analyzed. In the first year of the study, fox squirrel use of tree stands at the Pines course indicated no significant departure from expected (Table 10). The mixture of golf course property with private and condominium property at Site 2 created a more complex landscape than seen at Site 1 . Analysis of the 147 tracking locations within the 26.0 hectares of tree stands on the Royal Palm property showed that squirrels used the vegetation classes in proportion to their abundance on the landscape (X 2 =12.22, df=6, p=0.058). Pine plots account for 68% of the forest stands and contained 65% of the analyzed tracking points. Class 9, mixed natives with exotics, and class 3, cabbage palm dominated stands, had slightly higher than expected use. The 34.9 ha of condominium property and the private pine forest inside the back 9 of the course had only 34 tracking points. Nineteen of these were in the 1 .4 ha private pine forest that contained at least 3, well-stocked feeders. The 40. 1 ha of residential property accounted for only 17 tracking location, 10 at feeders.

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86 60% T Figure 32. Habitat preferences for all fox squirrels at the Pines course, Royal Poinciana. All tracking points within the mapped vegetation plots on the Pines course are included, n=164.

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87 ~ ^ CD > X <4H O .S c Q o IS -a 00 1 1 o c 53 o O OS 2 T XI fa r <2 (~| c/i cO 3 c/i v> .2 T3 O a .2 t/j — • ?s 0 ~ >> -g T3 0 3 1/1 CW 11 •§ -S -3 CO 1 O x =: <© 03 o 8 in o •j — 1 r \ I 8 [J— "3 ^ u c CO H 09 1) I O e a >> 6 J o Al II I a ad c 0 o u © II 8 8 5 o\ 2 ~ n© o on CO 00 vo 3 On < c 3 "* on wo V ~ x 2 £ on 3 < -a § s 1 ! — l/i cn ON ci 00 o 00 NO O — • — !} rm N oc Q. CO .1 ft "8 % ou U O I \0 — 00 OS CN M ^ t-« >4m — os <-< 00 Tt — 00 U1 1 in ci vo " 4 O -R < ^ • .a « .a o a O ii, H ffl B » 1^ 8 D o o 0. < CO < CS 2 X) C 0 0 0 0 0 0 «* OS oc Ss 00 CI rfs 00 0 vO Cs Tl•A ~~ 00 V J sD ON rfS r"* ON O 1— « rprt n >n n o \6 •4r sD ci I a

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88 Feeding Patterns Feeding data from Site 1 show seasonal shifts between native and non-native food sources (Table 1 1). Native foods, pine, cypress, and oaks, made up over 70% of the diet from August 1996 through January 1997. In March through July of 1996 and 1997, non-natives were the primary food source. February, June, and July of the 2 years showed a changing mixture of native and non-native sources. In 1996 squirrels began to feed on new, green pine cones in late June and early July. By August cones were mature and pine became the dominant food, with squirrels feeding alone or in small groups in or around select trees for several hours at a time. This also began the season of burying cones, in grass, litter and sand traps. Squirrels were observed burying in their core areas and at the edges of their home ranges. In December and January, cypress cones increased in importance, as pines decreased. Cypress constituted 20% or more of the diet for 7 months, July 1996 through January 1997. Despite the tent caterpillar infestation of 1997, cypress were again accounting for 29% of the diet in June and July of that year. Feeding on early pine cones began in June and July 1997. Oaks were not a regular food source in 1996 until September, when oak feeding, primarily live oak, accounted for 30% of the diet. Fox squirrels continued to feed lightly on oaks through April 1997. In both 1996 and 1997 fox squirrels took advantage of the bright red samara of the red maples for concentrated feeding in January and February. Late winter and spring feeding on natives was scattered. In April and May of both years squirrels fed on old pine cones, most often digging them up, but occasionally removing them from the tree. Cypress

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89 remained a minor element in the diet in all winter and spring months of 1996 and 1997, except February 1996. Fox squirrels fed on large hypogeous fungi from February through October. While they fed on both the mycelia and the fruiting bodies, the most readily observed behavior was the feeding on the large fruiting bodies. The peak season for observation of fungi feeding varied slightly, June and July in 1996 and May and June 1997. Fungi feeding was concentrated in patches that had deep litter layers, generally beneath pine and cabbage palm, but also in stands of cypress and pine. Of the 1 1 1 fungi feeding observations at 53 tracking points, 86% were in litter, 9% in grass, and 5% in trees. A squirrel might carry a fruiting body up a tree, where he would hang upside down and eat. In peak season, I recorded periods of concentrated feeding when a squirrel would eat the large caps (5-8 cm) of 4-5 fruiting bodies within 10 minutes. In May and June 1997 squirrels fed on concentrated patches of Mahogany webworm larva, Macalla thyrsialis (John Heppner, personal communication), buried in soil, grass or litter. They bit the tip from the cases and pulled the small caterpillars out with their teeth. During the same period they showed no interest in the tent caterpillars that rained to the ground from the infested cypress trees. In both 1996 and 1997, March through May were periods of concentrated feeding on non-natives. In March 1996 to June of that year, over 40% of recorded feeding was on bischofia {Bischofia javanica), an Asian species that forms clusters of small, dark berries. Squirrels initially fed in trees and then moved to ground feeding, where they gathered in groups of 2-8 to feed on the fallen fruit. Four large bischofia trees on the Cypress course

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90 drew squirrels in mid morning and late afternoon feeding groups. In 1997, levels of bischofia feeding were lower, with peaks in February through April in the 20-30% range. Bottlebrush trees, native to Australia, drew squirrels to feed on their spikes of scarlet flowers in March and April of both years. Silk oaks (Grevillea robusta), another Australian native, with their orange-fringed flowers, also attracted fox squirrels for flower feeding. In 1996 the peak feeding was late April and May, in 1997, late March and April. Queen palms, common throughout Royal Poinciana, produce a bright orange, aromatic, 2 cm oval fruit. A squirrel feeding on the fruit makes a distinctive, loud grating sound, whether hanging from clusters of new fruits or digging up a previously buried specimen, and so identification is easy. Queen palm feeding was high in February of both years and moderate in April and July of 1996 and May of 1997. Squirrels regularly buried queen palm fruits and most of the spring 1997 records are squirrels feeding on fruit they dug up. Heavy trimming of palms in the fall and winter of 1996-97, removed most of the available fresh crop. Regrowth and new fruits did not appear until summer of 1997. Ten large ficus trees (Ficus spp.) dotted the Cypress course and stands of large ficus were common at Hole in the Wall. These drew squirrels for feeding from January through August. A variety of species with staggered fruiting times produced the most concentrated feeding in May through July. Fruit of the tallow tree {Sapium sebiferum) and Java plum (Syzigium cumini) trees were feeding sites for short seasons. Tallow trees provided feeding in January and February and Java plum in June and July. Feeding on Java plum was high in July 1997 as the study ended, when squirrels gathered in 4 stands that produced aromatic crops of fruit on the Cypress course.

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91 The 1996 fall feeding on native species was the period with the lowest food diversity. In both years, diversity was higher in February through July. Diversity was noticeably higher in 1997, February to April and June, when a broader range of non-native species and a greater use of native species were seen in the feeding records. This difference was not only a result of changes in who was collared, but was reflected in the expanded diets of individuals collared both seasons. While 4 squirrels showed concentrated bischofia feeding in spring 1996 (8-23 observations), none of these repeated the pattern in 1997. All increased the number of species they used. Only 1 individual, subadult ROPO 27, showed concentrated bischofia feeding in spring 1997 and she regularly fed on the 2 large bischofia trees near her birth tree. Her mother, ROPO 2, fed on at least 10 species each spring. The feeding data at Site 2, covering January to July 1997, were aggregated due to low numbers (Table 12). Feeding on non-natives was dominant, as it was in the late winter and spring at Site 1 . Native foods included pines, larvae and mushrooms. Pine feeding, 16% of total, was primarily cones from previous seasons. Larvae of the same species as Site 1 and mushrooms accounted for an additional 16% of feeding records. Cypress were a minor element of the diet and not common at Site 2 or neighboring courses. Sixty-seven percent of the recorded feeding was on non-native sources, while 33% of the sightings were squirrels feeding at feeders. Ten feeders were available to fox squirrels, whether placed there for their use or for birds. Eight of the feeders were on Royal Palm property or adjoining residential property, 2 were on the west side of the neighboring Hibiscus course (Figure 25). Bottlebrush trees and queen palms provided feeding opportunities at Royal

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92 Palm, while Java plums and ficus were grouped in a small area of the Hibiscus course. Only ROPA 8 was seen begging, and that, successfully, at the public Hibiscus course.

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93 Table 12. Feeding patterns of fox squirrels at Site 2. Data include all feeding records at all locations used by the Site 2 collared squirrels from January 1997 through July 1997. All food items are listed. Non-natives Feeders Bottlebrush Java plum Ficus Silk oak Queen palm Begging 67% 33% 13% 9% 4% 2% 2% 2% Natives Pine, new & Larva Mushrooms Cypress old cones 33% 16% 9% 7% 2%

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DISCUSSION The goal of this study was to gain an understanding of golf courses as suitable habitat for Big Cypress fox squirrels. In the following discussion I will address this goal by integrating the data on fox squirrel ecology with that on course landscapes. I will first look at the characteristics of the 2 radio-collared populations in the high and low quality golf course landscapes. Then I will interpret the findings of the Landscape Evaluation Index. The two courses with collared populations will serve as reference points for the further interpretation of the broad range of golf course landscapes identified and ranked through the use of this index. Recommendations for design and management of course landscapes will be presented in the final chapter. Higher Quality Golf Course Landscape Site 1, Royal Poinciana Cypress Course, provided high quality fox squirrel habitat with few intrusions by traffic. The course rated 0.956 on the Landscape Evaluation Index, with large, moderately dense to dense tree stands of mixed natives dominated by pine, cypress and cabbage palms. Scattered exotics provided a majority of late winter and spring feeding sites. Numerous non-irrigated areas with pine litter ground cover provided centers for concentrated feeding on fungi. Mixed tree stands with large trees, bromeliads, and moderately trimmed palms provided a variety of protected nesting sites. 94

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95 As would be expected, adult females had smaller home ranges than males (Kantola and Humphrey 1990, Weigl et al. 1989). Mean home range size for adult females was 10.10 in 1996 and 16.40 in 1997. Home range means for adult males were 70.84 in 1996 and 90.91 in 1997, larger than previously reported in work on southeastern fox squirrels. Kantola and Humphrey's (1990) work with S. n. Shermani in north Florida showed female home ranges averaging 16.7 ha and male home ranges averaging 42.8 ha (harmonic mean). In North Carolina, Weigl et al. (1989) found an average female home range size for S. n. niger of 17.2 ha and an average male home range size of 26.6 ha (MCP). Site 1 adult male home ranges were 7 times larger than adult female home ranges in year 1 and 5.5 times larger in year 2, showing a much greater difference between the sexes than reported by previous workers in the southeast (Kantola and Humphrey 1990, Weigl et al. 1989). Males with a substantial portion of their home ranges in neighboring courses of generally lower quality had larger home ranges. The strong overlap seen in home ranges of both males and females at Site 1 appears to be unusual for southeastern fox squirrels (Kantola and Humphrey 1990, and Weigl et al 1989). Kantola and Humphrey found little overlap in female home ranges, while Weigl et al. (1989) note seasonal variation with no overlap of male home ranges in times of winter scarcity. Subadult home ranges for both years of the study averaged 18 .3 ha, not significantly different from adult females. Male 13 increased the mean size by 2.6 ha with at least 3 long day trips into the Pines course the month before he dispersed out of the club to the south. Though home range size for subadults remained consistent throughout the study, dispersal of subadults differed greatly between 1996 and 1997. Four of the 10 individuals,

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96 3 females and 1 male, collared as subadults in 1995-96 dispersed from the course in the spring and summer of 1996. Three of the 4 were frequently observed being chased by adults before they left the course. In 1997, none of the 7 squirrels collared as subadults dispersed before the end of July when collars were removed. None of the subadults was observed being chased by adults in the 1997 season. All continued to have a majority of their home range within that of their mothers and all were known to be 1 1-13 months of age when the study ended. The sample of dispersers was too small to allow for detection of a sex bias. Unbiased dispersal would be expected if dispersal is resource driven, as it appears to be in solitary, promiscuous squirrels such as red squirrels (Anderson 1989, Larsen and Boutin 1998) and gray squirrels (Thompson 1978). Interpretation of the differences between the 2 years is difficult. As Tappe and Guynn (1998) pointed out, little work pertaining to dispersal in fox squirrels has been reported. Much that is available is speculation. While some researchers have asserted that all fox squirrel juveniles disperse (Hansen et al. 1986, Koprowski 1991), that was not evident here. The relatively small samples of 1996 and 1997 did show that some fox squirrels do not disperse and that both males and females disperse. One explanation for the differences in dispersal is that Big Cypress fox squirrels may not disperse before one year of age, though previous research indicates that dispersal of fox squirrels would be expected before that age. In more northern fox squirrels dispersal of spring born individuals peaks in the fall before they are a year old (Allen 1943, Baumgartner 1943). Size and sexual condition at the time of capture indicated dispersing individuals in the current study were born in summer 1995 and so dispersed just before they were one year of age. So why did their counterparts in 1996 stay at home? Perhaps in

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97 years of lower resources more tropical subspecies such as Big Cypress fox squirrels can delay dispersal and reproduction until later in the season. Perhaps dispersal was driven by changes in resources or related changes in female reproductive cycles. Data indicate that resources were less favorable in 1997 than 1996, and there were differences in reproduction. In 1996, 4 of the 6 adult females had brood nests by the end of July. In summer 1997, only 1 of the 5 collared adult females was known to be attending a summer brood nest by the end of July. Interestingly, in spring and summer 1997, adult female ROPO 02 had a collared subadult from summer 1996 and offspring from early 1997 all using her core area, 3 generations, though none of the female offspring were reproductively active. If we consider that dispersal is resource related (Anderson 1989, Larsen and Boutin 1998), that agonistic behavior is the highest during periods of reproductive activity (Benson 1980), and that adult females are the most aggressive in motivating offspring to disperse (Adams 1984, Larsen and Boutin 1998), it would follow that adult females would motivate male and female subadults to leave in equal numbers in years of high resources when they are in a reproductive state. In years of lower resources females may not enter a reproductive state and may not encourage their offspring to leave, especially if resources are not critically low enough to threaten their own survival. Spring/summer 1996 was a time of high enough food to allow widespread reproduction and thus instigated aggressive behavior on the part of adult females toward subadults. Spring/summer 1997 was a year of lower food supply, maybe of an intermediate level, low enough to prevent reproduction, but not low enough to create chronic food stress in adults, thereby causing them to drive away subadults. This explanation is certainly tentative. The questions raised here do

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98 illustrate the need for research on dispersal and the driving mechanisms in a variety of environments. The dispersing subadults of 1996 did not fare well after they left Site 1 . Two of the 3 dispersing females were known dead on neighboring courses within the time of the study, 1 was hit by a vehicle, the other died after an injury of an unknown origin. Neither had successfully reproduced. The remaining 2 dispersers disappeared from neighboring courses within 2 months and were not sighted during 12 months of frequent searches. Courses to the west and north of Site 1 had LEI ratings of 0.44-0.30, while the course to the south, at which ROPO 14 died, had an LEI of 0.59. All these ratings were much lower than Site 1 . The area surrounding the cluster of 7 courses was dense development bordered by 2 of the busiest roadways in Collier County. Squirrels were not known outside of golf courses in the developed parts of the county. Successful dispersal away from the course cluster was highly unlikely. Non-dispersing individuals captured as subadults at Site 1 fared better than dispersers. In the 1995-1996 cohort of collared subadults, 50% of non-dispersers, 1 female and 2 males, remained at Site 1 throughout the study. Eighty-three percent of nondispersers remained on the course through December 1996. Two of the females successfully reproduced in summer 1996 and by the end of the study the 2 produced 7 young from the nest. One male of the 1996-96 subadult cohort was generally the lead male in mating chases by the end of the study. In 1997, 6 of the 7 collared subadults survived and remained on the course until collars were removed at the end of July. The 1 mortality was a subadult suffering from skin fungus. Two other subadults with the disease,

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99 1 a sibling of the dead squirrel, recovered with the spring/summer molt. Though none of the individuals reproduced, 1 female was the focus of a mating chase in July. The persistence and success of subadults that remained at Site 1 indicates it was providing a rich habitat. Dispersing subadults moved into less favorable environments. Site 1 appeared to be a source of fox squirrels for surrounding courses, which the 1996 data and the LEI ratings would indicate were acting as sinks (Pulliam 1988). Site 1 squirrel densities of 42.4 squirrels/km 2 (from high MNA of 26) to 49.8 squirrels/km 2 (from high Bailey estimate of 30.5) were higher than previous reports of fox squirrels in Florida. Estimates of 8.4 squirrels/km 2 and 38 squirrels/km 2 were reported by Humphrey et al. (1985) in south Florida and Moore (1957) in central Florida. In Georgia, at the Piedmont National Wildlife Refuge, Tappe et al. (1993) found fox squirrel densities of 15.3-17.7 squirrels/km 2 in mark and recapture studies. Fox squirrels at Site 1 had 2 breeding seasons, as reported in other southeastern fox squirrel populations (Moore 1957, Weigl et al. 1989). Contrary to previous work at more northern locations (Larson 1990, Weigl et al. 1989), reproduction was higher in the summer season than the 2 winter seasons. Litter size, as young from the nest, ranged from 1 to 4 with a mean of 2.4, within the reported range of 1 .6 to 3.0 (Larson 1990, Moore 1957, Weigl et al. 1989). While 4 of the 7 Site 1 adult females in the study produced 2 litters in the 3 breeding seasons studied, none produced 3 litters. Weigl et al. (1989) showed a strong correlation between food availability and female reproductive capacity. In the summer of 1996, 5 of 6 females produced young from the nest and raised them into the fall. The smaller home ranges in that year and the concentrated feeding patterns of

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100 spring through fall of 1996 indicate a rich food supply, probably resulting from the high rains of fall 1995. Frequent squirrel feeding on fungi was also reported by Weigl et al. (1989) in North Carolina. Presence of mycorrhizal hypogeous fungi and associated bacteria form mutualistic relationships with a variety of tree species, including pines (Li et al. 1986, Slankis 1973).The fungi are nutritionally beneficial to fox squirrels who then disperse fungal spores as they defecate in their wide travels (Maser et al. 1978, Trappe and Maser 1977). Maintenance of the fungi-rich litter areas within Site 1 preserved a food source for fox squirrels and probably provided benefit to a tree species also of primary importance to the squirrels. Weigl et al. (1989) believed the fitness of squirrels, trees and fungi benefited from this relationship and that it may be coevolutionary in nature. High densities, high reproduction, and high overlap of home ranges suggested a food supply that was strong though obviously variable. Occupancy of Site 1 by 5 or 6 adult females maintaining fairly constant home ranges throughout the 20-month study indicated the adult female density was at a maximum. The course was able to retain subadults in both years. In 1996, 60% of the collared subadults remained on the course, with 50% of the non-dispersers surviving through the end of the study. In 1997, 6 of the 7 collared subadults remained alive on the course until the end of the study. Site 1 and its companion course, Pines, together had 40-50 squirrels in residence, though it must be noted that at least half of the adult males used neighboring clubs. The Site 1 population appeared stable. Surplus subadults dispersed to neighboring courses where mortality undoubtedly was higher.

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101 Royal Poinciana and its buffer courses to the west provided the most productive and stable golf course site for fox squirrels within the currently developed landscape in western Collier county and the most stable in this study. Drastic changes in the landscapes of Poinciana or the 2 neighboring courses most frequently used by males are not anticipated in the next 5-10 years. On the other hand, these landscapes are managed. The composition and maintenance of trees, tree stands and ground cover are subject to change at the will of the managers. East and south of Site 1, the condition of neighboring properties will change in the coming decade, with anticipated development of the large pine forest south of Poinciana and current development of the lots east of the club. Such conversion will affect sources and movement of squirrels in and out of Poinciana and could bring more domestic predators, especially cats, to the borders of the courses. Lower Quality Golf Course Landscape Site 2, Royal Palm Country Club, just one year older than Site 1, contained lower quality fox squirrel habitat surrounded by a developed landscape and a seasonally heavy flow of vehicle traffic. The club rated 0.719 on the Landscape Evaluation Index with moderately dense to open pine-dominated tree stands on club property and neighboring condominium land. There was a lower diversity of tree species, both native and nonnative, and fewer and more widely scattered pine litter areas than at Site 1 . In the absence of oaks and maples, and with lower numbers of large fungi feeding areas or fruiting exotics, squirrels fed heavily at feeders in the spring and summer. The 2 adult females at Site 2 had home ranges of 13 .06 ha and 30.57 ha. The difference probably resulted from the former having a regularly supplied feeder in the

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102 center of her home range and the latter having only an occasionally stocked one at the edge of her home range. The home range of the first female was similar to those at Site 1 and to previous studies, while the home range of the second female was larger than any other female in this study and the averages reported in other studies (Kantola and Humphrey 1990, Weigl et al. 1989). Male home ranges of 136.1 ha and 303.80 ha were larger than others in the study and larger than means reported by Kantola and Humphrey (1990) or Weigl et al. (1989). All adult and subadult home ranges at Site 2, except the 1 adult female with a well-stocked feeder, were larger than comparable individuals at Site 1 and larger than reported means in previous studies. Adult female home ranges did not overlap or touch as they did at Site 1 . While the 4 collared adults lived and persisted on the course through the 7 months of tracking, none of the collared subadults remained. Two of the 4 subadults were known dead by June 1997, the third had a habit of feeding in roadways and disappeared early, and the forth dispersed to a very low quality habitat in an adjoining course and was using a feeder near a major highway. One of the subadults died from extreme emaciation while suffering a severe skin fungus, the other was killed by a car on Augusta Boulevard. Vehicle accidents at the study sites involving automobiles, course equipment and carts, were a known source of mortality for 6 collared and uncollared squirrels . Weigl et al. (1989) found that automobile traffic was a major cause of fox squirrel mortality in their 8year study in rural areas of the North Carolina coastal plain. Neither adult female reproduced during the study, though 1 appeared to be tending a brood nest as the study ended. Both males were observed with females at Site 2 and the 2 neighboring courses.

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103 Fox squirrel densities at Site 2 were extremely low, 6.3/km 2 (MNA of 9) and 8.2/km 2 (high Bailey estimate of 11.7). These were similar to the estimates of Humphrey et al.(1985), though much lower than reported by Moore (1957) or Tappe et al. (1993). They were 84% lower than the fox squirrel densities at Site 1 . At Royal Palm, low population density, large home ranges, and poor subadult persistence indicate an unstable population. A small number of adults maintained themselves and reproduced in times of higher food supply, but young had a difficult time surviving to adulthood. A more open, less diverse food supply required fox squirrels to make larger movements to feed and mate, and so expose themselves to the hazards of automobile traffic and the stresses of travel and food search. Such large home ranges and movement through traffic were especially difficult for subadults. Royal Palm was a course with marginal fox squirrel habitat and more development yet to come. Completion of housing along the fairways will eliminate the remaining treecovered vacant lots. Growth of adjacent developments to the east and north will increase traffic on Augusta Boulevard and connecting roadways. As with Site 1, the quality of course vegetation is dependent on management. It could be improved with the addition of a variety of native trees such as cypress, oaks, and maples, with added pine litter areas, and with non-native food sources such as spring-fruiting trees. While feeders were obviously a vital part of squirrel diets at Site 2, reliance on feeders puts squirrels at the mercy of suppliers and perhaps increases the risk of exposure to contagious disease. The course adjoining Royal Palm to the west, used by adult males and at least 2 dispersing subadults in the study, had very low quality habitat (LEI 0.28) and would not support fox squirrels for long without the resources of Royal Palm.

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104 As we shall see in the next section of the discussion, Site 1 was an unusual golf course in southwest Florida, both in its rich landscape without residential development and in its high numbers of fox squirrels. Site 2, on the other hand, was more common. It was a mix of the favorable and unfavorable landscape features. It had the levels of development and traffic seen in many courses of level 3 and 4, and though it was not isolated, the 2 neighboring courses furnished very low quality fox squirrel habitat. It was superior to most courses in the presence of relatively high quality pine stands, the open understory of all tree stands, and the occurrence of scattered pine litter areas. Landscape Evaluation The study revealed a wide variety of golf course habitat types and course configurations in Lee and Collier counties. Landscapes surrounding these courses, which are critical to fox squirrel movements, ranged from highly developed with heavy vehicle traffic to more rural sites surrounded by mixed agricultural and forest stands. The ability of golf courses to support fox squirrels differed greatly. Six of the courses with an LEI of 0.90 or above had high levels of fox squirrels and make up 3 clubs, each of 36 holes (Fig. 33, Appendix B). These 3 clubs had the highest potential for supporting fox squirrels in a developed landscape. The 6 courses with an LEI of 0.90 and above and with high levels of fox squirrels, including Site 1, were characterized by . • large contiguous areas, over 120 hectares, with no housing or automobile traffic • residential development absent or only on the perimeter of each course or the entire club • adjoining golf courses on at least 2 sides of the club • undeveloped forest on at least 1 side • lack of busy roadways around the course

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105 • easy movement of fox squirrels from 1 course to another, often aided by large trees across canals or smaller streets • large, open, pine-rich tree stands, with cypress, palms and a variety of native tree species • few or no areas of forest with heavy understory growth • high-quality nesting sites in minimally trimmed cabbage palms, bromeliads, or large trees, often pine and cypress • spring and early summer food supplies available from diverse native species, fungi rich Utter areas, non-native tree species, or artificial food sources. Four of these 6 high level courses will have further housing development along the fairways and all will have increased development around their boundaries within the next 10 years. Except for Site 1 and its companion Pines course, squirrels fed from feeders or begged from golfers. The future of fox squirrels at even these 6 high level courses will depend on the maintenance of high quality tree stands through understory clearing, and planting and replacement of native trees, especially pines and cypress. Changes in surrounding landscapes and food supplies offered at feeders will also affect squirrel survival. Given the lack of development at Site 1 and the relative protection of the surrounding buffering courses it appeared to offer a relatively stable, high quality environment for fox squirrels, though it is certainly suburban and will eventually become more isolated. The other high level courses will undergo much more change with stronger potential for declining habitat. The 1 course with an LEI of 0.90 and moderate levels of fox squirrels was also part of a 36 hole course, but its companion course had an LEI of 0.71, with high traffic and more intense development in the future. This 0.90 LEI course does not have a strong potential for continued support of fox squirrel populations.

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106 j ^ * * I J > o ta « "2 i -3 00 > i TO 511 a — >„ V CO CM |3.\aq uojiKindoj pjjinbs .1

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107 The 7 courses with an LEI over 0.90 obviously provided habitat superior to any other courses in the study and each had a strong combination of favorable landscape attributes. The remaining 53 courses had LEI ratings of 0.76 or lower (Fig. 33, Appendix B). Each of these courses had one or more strong negative elements in their landscapes. Features of isolation, course configuration, low quality vegetation, and heavy understory, combined to decrease the ability of these courses to foster fox squirrel populations. Course isolation within a heavily developed landscape cannot be mitigated in most cases. This is especially true if the course is small and contains few or scattered tree stands. A group of 9 older courses with low LEI ratings is circled in the lower portion of Figure 33. Each of these courses was the only course in an 18 hole club. The courses may have experienced loss of trees with long-term development. They frequently had few remaining tree stands and low levels of native tree species, especially pines. They were surrounded by residential and commercial development, though some were adjacent to similar courses. At first glance they appeared to offer fox squirrel habitat, but in fact, they provided insufficient food and nesting resources for this relatively large squirrel species. Complex course configuration, as presented previously in Figure 8, levels B and C, were a common landscape element that created precarious habitat for fox squirrels. Unlike the higher LEI courses that had no development or only perimeter development, 39 of the 53 courses (74%) with LEI below 0.75 had intensified or connected perimeter patterns, or radiating interior development. Such development configurations divided the course and the bordering rough areas into small patches. These patches may or may not have contained high quality tree stands, but these divided landscapes required squirrels to move though a maze of housing and streets in search of food, mates, and nesting sites. For

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108 animals with home ranges of 10 to 100 hectares and more, the increased and complicated fragmentation of an already fragmented landscape becomes even more precarious and stressful, as resources become more widely spread (Collinge 1998). Development of courses with complex configurations creates a landscape that is ultimately highly unfavorable to fox squirrels. Deceptively, initial and temporary stages of development may actually improve habitat for fox squirrels. The early development may increase edges and open the forests understory, thus creating the habitat fox squirrels prefer. Eighteen of the level 3 courses (72%) are at this stage. Unfortunately, as courses age and development continues, construction removes tree stands and corridors required by fox squirrels. Vehicle traffic within and around a course increases and the habitat becomes less productive and more stressful for fox squirrels. The result, as seen at Site 2, is that fox squirrel home ranges must become exceptionally large to reach scattered resources. Constant travel in a developed landscape from one feeding patch to another is stressful and hazardous. This situation is especially difficult for younger squirrels. As these small populations of widely scattered individuals become more isolated, they become more susceptible to stochastic events or dying out in years of low food production. Low quality vegetation stands were common in the golf course landscapes. Few courses had a high diversity of native species or large pine stands, few had large areas of pine litter to support growth of fungi, and few had the older, large trees that offer ideal nesting sites. The first component of the LEI indicates the quality of the vegetation at a course. The 6 high level courses each have a rating of 0.9 and higher in this component. Of the remaining 54 courses, only 4 are above 0.80 in this component, while 30 of the courses, 50% of those in the study, have ratings below 0.6.

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109 Heavy understory, with the dense growth of vines and shrubs in tree stands, is a landscape element that renders habitat unsuitable to fox squirrels. It was a common and difficult problem in unmanaged set-aside areas. Eighteen courses in the study (30%) had varying amounts of heavy understory in their landscapes. For 16 of the 18 courses it was just one impediment to fox squirrel presence. Four of the 16 courses are circled in Figure 33. They will continue to be developed and have complex course configurations with highly fragmented habitat. Clearing of the understory in these courses will not create quality fox squirrel habitat The four prominent elements that affect the quality of fox squirrel habitat on golf courses vary in their ability to be changed through good management. Two of these, course isolation and course configuration are critical elements affecting squirrel movements and the availability of resources. They must be addressed prior to development. The latter two, the presence of heavy understory, and the composition and density of tree stands, can be mitigated to some degree on an existing course, though they will not nullify the impacts of isolation, heavy development, and poor course configuration in the long run. Mitigation of heavy understory growth in set-aside areas is especially difficult and expensive. The improvement of the tree stands and ground cover should be encouraged on courses that currently have fox squirrels in residence or in the adjacent lands. While all level 3 and 4 courses are candidates for vegetation improvement, the 1 1 courses that have high quality courses for neighbors should be strongly encouraged to undertake habitat improvement for fox squirrels (Fig. 33). Work to increase and diversify native tree species, to create clear understory, to increase the number of moderately trimmed palms, to

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110 increase spring food sources, and to increase areas of pine litter ground cover will improve habitat. In addition to this larger group of level 3 and 4 courses, two relatively new, nonresidential courses east of 1-75 warrant special attention for habitat improvement efforts. The 2 courses, each 18 holes at the time of the study, were part of clubs that will become 36 holes or larger. The courses contained large stands of pines, cypress, palms, and associated native tree species. They were surrounded by undeveloped forests, large-lot residential areas, and agriculture. They were less than 3 km apart. At the time of the study both courses had heavy understory growth and the resulting low LEI ratings along with low numbers of fox squirrels. Habitat improvement through understory clearing would surely increase the potential for these clubs as fox squirrel habitat. Their position in a less developed landscape and their non-residential status gives them a unique opportunity to provide habitat for Big Cypress fox squirrels. Summary Will golf courses provide habitat for Big Cypress fox squirrels in rapidly developing southwest Florida where human populations are expected to double by 2020? As noted earlier, even the 6 courses with high levels of squirrels do not all have a strong potential as future fox squirrel habitat. Of these 6, Site 1 and its companion course at Royal Poinciana, offer the most favorable and most secure habitat for fox squirrels over the next 2 decades. The other 4, as residential courses, will continue to be developed and will have greater changes within and around their boundaries. Their potential will decline.

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Ill The remaining 54 courses will not provide good, long-term resources for fox squirrels. Certainly, all of the 23 courses in levels 1 and 2, have more than one strongly negative landscape element and most do not have the potential to support fox squirrels even with mitigation of vegetation. Most of the 31 courses at level 3 and 4 will not provide good quality habitat for the long-term due to unalterable planning and design patterns. Many of these are relatively new courses where time and development will continue to degrade the remaining tree stands and unmanaged set-aside areas. This study demonstrated that even in extremely high quality habitat, Big Cypress fox squirrels require large tracts of land for daily and seasonal movements and even larger ones to allow for dispersal of subadults. Few courses or groups of courses offer safe and stable habitat in large enough tracts to endure the upcoming intensity of development, especially in the western sections of Lee and Collier counties. The few that do must maintain open, diverse, pine-rich forested areas, preferably with substantial areas of pine litter ground cover. Maintenance of such a landscape is labor-intensive and expensive. Fewer than 5 of the 48 clubs examined in this study are capable of providing the habitat required to maintain golf course populations through the intensive development expected between now and 2020.

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MANAGEMENT RECOMMENDATIONS Management for quality Big Cypress fox squirrel habitat must start at the landscape level. Placement and configuration of courses and the development that accompanies them is critical. Because fox squirrels use large areas, up to 150 hectares, for daily and seasonal movements, they require large contiguous areas of suitable habitat free from vehicle traffic and dense development. They must be able to move from one club to another as few single courses will provide adequate habitat. Developments should be connected by open forested corridors. Movement across 2 lane roadways can be facilitated by maintaining large over-hanging trees. Squirrels have adapted to using wooden walkways through wetlands at several southwest Florida courses. This indicates they would be able to do so to cross other obstructions, perhaps even busy roadways separating 2 courses or clusters of courses. A course or club must be designed to contain large contiguous areas, 120 hectares per club, of green space with large, open stands of native trees. This should be accomplished by concentrating the development either around the outside edges and leaving the central area of the course as green space or by concentrating development in the center, with only 1 roadway into the developed center. This creates a large circle of green space free of vehicle traffic. Adjoining courses with this central development could provide large areas for wildlife. 112

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113 In more recently developed courses, the design and planning phase has included much attention to preserving lands within the developed areas that have been designated as critical wetlands or drier pine habitat for gopher tortoises. This often appears to preserve large areas of habitat suitable for fox squirrels. In truth, the habitats do not remain suitable for many of the species they are designed to conserve, and they generally become unsuitable for fox squirrels. Areas set aside as native habitat are allowed to become clogged with invasive vines and a heavy understory of native and exotic shrubs and small trees. The resulting vine-infested forests often become barriers to wildlife movement, instead of habitat and corridors. Without proper training in management of wild habitats or the funding to carry out the required tasks of burning or hand clearing, managers cannot preserve these habitats. If such patches are to be preserved within private property, funding for maintenance and training must be provided and regular checks must be made to see that landowners comply with management plans. The management and usefulness of these areas require examination. Are they the best way to preserve wildlife habitat? Are they proving beneficial to the species they are intended to protect? If they are to be maintained, what programs are required to insure they fulfill their expected roles in the landscape? Big Cypress fox squirrels have shown a strong dependence on pines and cypress, using them for food, nesting, and resting areas. Preservation and planting of pines and cypress should be strongly encouraged. Golf courses are currently moving away from full coverage irrigation and this will hopefully allow more native pines to persist in developed landscapes. In addition to pine and cypress, planting a diversity of native trees, including oaks, maples, cabbage palms, bays, and hollies should be encouraged. The current study

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114 showed a heavy use of spring-fruiting non-native tree species by fox squirrels. Though the planting of non-natives is against the policy of some organizations promoting more natural golf course landscapes, presence of these non-native species provides year-round food in these restricted habitats. Managers should create diverse stands of trees that provide a range of food sources within a small space. This forms a richer, more resilient food source and provides the diverse environment fox squirrels prefer for brood nests. Course designers and managers must have information on native plant sources, and plants must be available at competitive prices before native plantings will become common. On completed courses, correct management of existing tree stands is crucial. Fox squirrels require open tree stands for movement and frequent ground feeding. Forested areas must have open understories, free of dense shrubs, vines and tall grasses. This can be accomplished by hand clearing or light burning. At present, it is extremely difficult to obtain burning permits for golf courses, despite the fact that ready irrigation systems provide excellent protection against uncontrolled burns. The smoke created by such a burn is undoubtedly less harmful than most of the chemicals used in hand clearing. Progress along these lines, and studies addressing the efficacy of burning golf course roughs, would help the more forward thinking managers who would like to promote burning as a management tool for larger forested stands within their courses. Cabbage palms provide fox squirrels with high quality nesting sites, food, and sheltered resting areas. Current golf course management practices frequently involve extreme trimming of palms. This removes all the fruiting stalks and the lower leaves. What remains is barely a tree and is not habitat for the range of wildlife frequently seeking protection from sun and storms under the layered leaves and long leaf bases. Moderation

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115 in trimming is one of the easiest and least expensive management techniques benefiting fox squirrels. It should be encouraged. The presence of pine litter ground layers in pine stands is an important management technique that promotes squirrel feeding on fungi. The fungi have been shown to be beneficial to pines and provide a needed food source in early summer months. The litter layers can readily reduce maintenance and remove the need for irrigation of grass in a stand of native trees. If nesting sites in pines, large bromeliads, cabbage palms, and cypress are in short supply, managers may wish to supplement with nest boxes. Wood duck boxes are the proper size and should be placed at least 5 meters up the trunk of a fairly large tree. Such boxes will provide shelter in the few times of extreme cold weather and in driving rain and wind storms. They may also be used for brood nests. Education is critical for management of fox squirrels on golf courses. Managers must be educated on methods to create and maintain favorable habitat. Members should also be educated. Squirrels are easily killed on courses by cart drivers who are speeding along and looking for golf balls instead of squirrels. Members should at least be encouraged to look out for squirrels, on the course and on the roadways into the course. Members must also be educated not to feed squirrels. If squirrels are not fed by people they will not be attracted to people or carts. They will avoid carts and not hang around cart paths at tees and greens waiting for food. They will be less likely to be killed by speeding carts or angry golfers who have just lost a muffin to a sneaky squirrel. If course members or managers have the desire to feed fox squirrels, this should be done in an isolated area away from cart or automobile traffic. Food should be spread on

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116 the ground, not in small feeders that require squirrels to climb around on the same small area. Such high concentration and repeated rubbing of fur and feet on a feeder creates an ideal vector for contagious diseases, especially skin fungus. Food should be natural, not processed human food. Squirrels benefit from nuts, berries, and grains. Trimmed fruits from palms may also be added to the feeding site. Both club members and fox squirrels will benefit if education programs share something of the lives of fox squirrels. Many golf course members in Florida are not familiar with our native wildlife and plants. A little education may go a long way in helping them to understand and appreciate these unique and beautiful fox squirrels and the larger natural heritage of Florida.

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APPENDIX A TREE SPECIES ON STUDY SITES Common Name Family Species Site 1 Cypress Site 2 Royal Palm Royal Poinciana Pines Red maple Holly Schefflera Norfolk Island Pine Queen palm Fishtail palm Royal palm Cabbage palm Jacaranda Trumpet Tree Toog Tree Austrailian Pine Black olive Southern Red Cedar Tallow tree Earleaf acacia Rosewood Poincianna Copper pod Pongam Laurel oak Live oak Aceraceae Aquifoliaceae Araliaceae Araucariaceae Arecaceae Arecaceae Arecaceae Arecaceae Bignoniaceae Bignoniaceae Bischofiaceae Casuarinaceae Combretaceae Cupressaceae Euphorbiaceae Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae Fagaceae Fagaceae Acer rubrum x Ilex opaca x x Schefflera x actinophylla Araucaria x heterophyla Arecastrum x x romanzoffianum Caryota mitis x Roy stoma spp. x x Sabal palmetto x x Jacaranda x x mimosffolia Tabebuia argentea x Bischofia javanica x Casuarina x cunninghamiana Bucida buceras x x Juniper silicicola Sapium sebiferum x x Acacia x x auriculiformis Dahlbergia sissoo x x Delonix regia x Peltaphorum x pterocarpum Pongamia pinnata Quercus laurifolia x x Quercus x x virginianum x x x x x x x x x x x x X X X X 117

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118 Red bay Lauraceae Avocado Lauraceie Mahogany Meliaceae Wild Tamannd Mimosaceae Fig Moraceae Eucalyptus Myrtaceae Eugenia Myrtaceae Java plum Myrtaceae Bottle-brush Myrtacece White Ash Oleaceae Screw Pine Pandanaceae Slash pine "TV * Pinaceae Silk oak Proteaceae Orange Rutaceae Grapefruit Rutaceae Pond cypress Taxodiaceae Bald cypress Taxodiaceae Per sea borbonia Persea americana Swietenia mahogani Lysiloma latisiliquum Ficus spp. Eucalyptus sp. Eugenia sp. Syzigium cumini Callistemon rigidus Fraxinus americana Pandanus utilis Pinus elliottii var. densa Grevillea robusta Citrus sinensis Citrus x paradisi Taxodium ascendens Taxodium distichum

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APPENDIX B GOLF COURSES, SQUIRREL LEVELS, AND LEI RANKINGS Club Course Course # Squirrel level LEI l^Udll l^ICCK f~Vi i ail ircp 30 J 0.987 V^Udll V^ICCK. i r*oolr r»r\i ire* 1 VylCCK vUUi jC 2Q 5 0.987 ivoydi roint/idnd 46 5 0.979 Royal Poinciana cypress uourse to c j Fiddlesticks Long iviean c J Fiddlesticks ripers L-nanenge '{I c J n 004 Imperial imperial nasi 4J A t 0 000 Wyndemere Lrreen ana wnite 1*5 jL J 0 7*»1 nagie tuage Zo a 0 74S ivoyai r aim JZ 4 0 719 Imperial • Imperial West 4 J 0 711 V) . / 1-5 Forest Bear JO 0 713 v^uaii west preserve course 1 ? 1 z j 0 687 Lely Country Club Flamingo Island 1 1 1 / i 3 u.o m Lely Country Club Classics Course 1 1 1 1 J U.Oj j Forest Bobcat 1 A 1 D -J J n ^i o u.o iz Foxfire old lo, red 3!) s U.OUj Pelican Nest Crator otbeminoie zj 5 u.ouu Bonita Bay Marsh Course 6 1 U. jy!) Bears Paw 41 3 f\ coo U. joy Wyndemere Gold course 1 1 j l i 3 0 V.joj Old Hickory 13 4 0.571 Name withheld 5 3 0.555 Bonita Bay Bay Island 7 1 0.554 Countryside GC 22 3 0.554 Wildcat Run 23 3 0.541 Foxfire new 9 3 2 0.530 Royal Wood 19 3 0.525 Glades Palmetto 44 3 0.522 Spanish Wells North 9, New 9 2 2 0.516 Pelican Marsh Marsh Course 8 1 0.491 Glades Pines 53 2 0.487 Spanish Wells east and west 9 39 3 0.475 119

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120 Club Course Course # Squirrel level LEI Vines 24 1 0.473 Olde Florida 9 2 0.470 Wilderness 42 3 0.443 Quail West Lakes 4 1 0.388 Eastwood 40 2 0.365 Colliers Reserve 10 3 0.365 Marriott Club 15 2 0.363 Audubon CC 18 3 0.361 Hole in the Wall 58 4 0.354 Vineyards North 21 2 0.354 Embassy Woods 14 2 0.354 CC of Naples 55 4 0.343 Vineyards South 20 3 0.309 Bonita Bay Cypress Course 1 3 0.309 Quail Run 50 3 0.296 Hibiscus 56 3 0.283 Palm River 38 3 0.281 Cross Creek 27 2 0.275 Marco Shores 49 1 0.243 Myerlee 47 1 0.235 Windstar 26 1 0.214 Ft. Myers GC 60 1 0.204 Cypress Lake 54 1 0.200 Naples Beach 59 0.196 Moorings 57 0.196 Whiskey Creek 51 0.184 Club at Pelican Bay 37 0.000

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APPENDIX C HOME RANGE DATA FOR SITES 1 AND 2

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LIST OF REFERENCES Adams, C. E. 1984. Diversity in fox squirrel spatial relationships and activity rhythms. Texas Journal of Science, 36: 197-205. Allen, D. L. 1943. Michigan fox squirrel management. Michigan Department of Conversation Game Division Publication. 100: 1-404. Anderson, P. K. 1989. Dispersal in rodents: a resident fitness hypothesis. Special Publication, the American Society of Mammalogists. 9: 1-141. Bailey, N. T. J. 1952. Improvements in the interpretation of recapture data. Journal of Animal Ecology. 21:120-127. Balough, J.C. and W.J. Walker, (editors) 1992. Golf Course Management and Construction. Environmental Issues. Lewis Publishers, Boca Raton. 951pp. Baumgartner, L. L. 1943. Fox squirrels in Ohio. The Journal of Wildlife Management. 26: 208-211. Bender, Louis C, Gary J. Roloff, and Jonathan B. Haufler. 1996. Evaluating confidence intervals for habitat suitability models. Wildlife Society Bulletin. 24(2): 347-3 52. Benson, B. N. 1980. Dominance relationships, mating behaviour and scent marking in fox squirrels (Sciurus niger). Mammalial 44: 143-160. Brooks, Robert P. 1997. Improving habitat suitability index models. Wildlife Society Bulletin. 25(1): 163-167. Brown, R.B., E. L. Stone, and V. W. Carlisle. 1990. Soils. Pages 35-69 in Ronald L. Myers and John J. Ewel, eds. Ecosystems of Florida. University of Central Florida Press, Orlando. Carr, M. H., J.D. Lambert, and P.D. Zwick. 1994. Mapping of continuous biological corridor potential in Central America. Final Report, Paseo Pantera, University of Florida. Chen, E., and J.F. Gerber. 1990. Climate. Pages 1 1-34 in Ronald L. Myers and John J. Ewel, eds. Ecosystems of Florida. University of Central Florida Press, Orlando. 125

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126 Collier County Community Development and Environmental Services Division. 1996. Collier County 1996, Demographic and Economic Profile. Naples, Florida. Collinge, Sharon K. 1998. Spatial arrangement of habitat patches and corridors: clues from ecological field experiments. Landscape and Urban Planning. 42: 157-168. Cresswell, W., and G. Smith. 1992. The effects of temporally autocorrelated data on methods of home range analysis. Pages 272-284 in Imants G. Priede and Susan M. Swift, eds. Wildlife telemetry: Remote monitoring and tracking of animals. Horwood, New York. Dodson, Ronald G. 1990. Audobon cooperative sanctuaries for golf course management. USA Green Section Record. 28:14-16. Dodson, Ronald G. 1994. On course with nature, but can you prove it?. USA Green Section Record. 32:15. Edmondson, Jolee. 1987. Hazards of the game. Audobon. 89(6):24-28. European Golf Association Ecology Unit. 1995. An environmental strategy for golf in Europe. Information Press, Oxford. 41pp. Fitzgibbon, Clare D. 1993. The distribution of grey squirrel dreys in farm woodland: the influence of wood area, isolation and management. Journal of Applied Ecology. 30:736-742. Forman, RT.T. 1995. Land: the ecology of landscapes and regions. Cambridge University Press. Fowler, J., and Lou Cohen. 1990. Practical statistics for field biology. Wiley and Sons, New York. 227pp. Foy, John H. 1989. Golf courses and the environment: what's the future? USA Green Section Record. 27:1-4. Grigg, G. T. 1990. Seeking a fresh vision of environmental responsibility. Golf Course Management 58(9):38-46. Hall, E. R. 1981. The mammals of North America. John Wiley and Sons, New York 1: 600pp. Hansen, L. P., C. M. Nixon, and S. P. Havera. 1986. Recapture rates and length of residence in an unexploited fox squirrel population. The American Midland Naturalist. 115:209-215.

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127 Humphrey, S. R , and P. G. R Jodice. 1992. Big Cypress fox squirrels {Sciurus niger avicennia). Pages 224-233 in S. R. Humphrey, ed. Rare and endangered biota of Florida. Vol. 1: Mammals. University Press of Florida, Gainesville. 392pp. Humphrey, S. R., J. F. Eisenberg, and R. Franz. 1985. Possibilities for restoring wildlife of a longleaf pine savanna in an abandoned citrus grove. Wildlife Society Bulletin, 13:487-496. Jodice, P. G. R. 1990. Ecology and translocation of urban populations of Big cypress fox squirrels (Sciurus niger avicennia). M. S. thesis, University of Florida, Gainesville. 89pp. Jodice, P. G. R , and S. R. Humphrey. 1992. Activity and diet of urban Big Cypress fox squirrels. J. Widl. Manage. 56(4): 685-692. Jodice, P. G. R, and S. R. Humphrey. 1993. Activity and diet of urban Big Cypress fox squirrels: a reply. J. Widl. Manage. 57(4)L 930-933. Kantola, A.T., and S. R. Humphrey. 1990. Home range and mast crops of Sherman's fox squirrel in Florida. J. Mammal. 71:41 1-419. Kenward, R. 1992. Quantity versus quality: programmed collection and analysis of radiotracking data. Pages 231-247 in Imants G. Priede and Susan M. Swift, eds. Wildlife telemetry: Remote monitoring and tracking of animals. Horwood. New York. Kie, John, J. Baldwin, and C. Evans. 1996. CALHOME: a program for estimating animal home ranges. Wildlife Society Bulletin, 24 (2):342-344. Koprowski, John L. 1991, The evolution of sociality in tree squirrels: the comparative behavioral ecology of fox squirrels and eastern gray squirrels. Ph.D. dissert., University of Kansas, Lawrence, 1 16pp. Krebs, C. J. 1999. Ecological methodology. Benjamin Cummings, Menlo Park, California. Larsen, Karl W., and Stan Boutin. 1998. Sex-unbiased philopatry in the North American red squirrel: (Tamiasciurus huasonicus). Pages 21-32. w Michael A. Steele, Joseph R. Merritt, and David A. Zegers, eds. Ecology and evolutionary biology of tree squirrels. Virginia Museum of Natural History, Special Publication Number 6. Larson, B. J. 1990. Habitat utilization, population dynamics and long-term viability in an insular population of Delmarva fox squirrels (Sciurus niger cinereus). M. S. thesis, University of Virginia, 87pp.

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128 Lee County Department of Community Development. 1998. The Lee Plan, 1998 Codification. Fort Myers, Florida. Leuzinger, P.V. 1994. A new attitude: Audobon, our golf course, and the community. USA Green Section Record. 32:2-5. Li, C. Y., C. Maser, and B. A. Caldwell. 1986. Role of rodents in forest nitrogen fixation: another aspect of mammal-mycorrhizal fungus-tree mutualism. Great Basin Naturalist 46:411-414. MacArthur, R. H. 1972. Geographical ecology. Harper and Row, New York, New York. Maehr, D. S. 1993. Activity and diet of an urban population of Big Cypress fox squirrels: a comment. J. Wildl. Manage. 5 7(4): 929-930. Maser, C, J. M. Trappe, and R. A. Nussbaum. 1978. Fungal-small mammal interrelations with emphasis on Oregon coniferous forests. Ecology 59:799-809. Mazzotti, Frank J., and Carol S. Morganstern. 1997. A scientific framework for managing urban natural areas. Landscape and Urban Planning 38:171-181. Mech, L. D. 1983. Handbook of Animal Radio-tracking. University of Minnesota Press, Minneapolis. Mech, L . D. 1998. Estimated cost of maintaining a recovered wolf population in agricultural regions of Minnesota. Wildlife Society Bulletin 26(4): 8 17-822. Mladenoff, David J., T.A. Sickley and A. P. Wydeven. 1999. Predicting gray wolf landscape recolonization: logistic regression models vs. new field data. Ecological Applications 9(l):37-44. Moore, J. C. 1954. Fox squirrel receptionists. Everglades Natural History. 2(3): 153-160. Moore, J. C. 1956. Variation in the fox squirrel in Florida. Am. Midland Nat. 55:41-65. Moore, J. C. 1957. The natural history of the fox squirrel, Sciurus niger shermani. Bull. Am. Mus. Nat. Hist. 113:1-71. Moul, I.E. and John Elliott. 1994. The bird community found on golf courses in British Columbia. Northwestern Naturalist 75:88-96. National Oceanic and Atmospheric Administration (NOAA). 1995-1997. Climatological Data, Florida. Vol. 99-101, nos. 1-13.

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129 Pearlstine, Leonard G., Laura A. Brandt, Frank J. Mazzotti, and Wiley M. Kitchens. 1997. Fragmentation of pine flatwood and marsh communities converted for ranching and citrus. Landscape and Urban Planning. 38: 159-169. Pulliam, H. R. 1988. Sources, sinks, and population regulation. American Naturalist 132: 652-661. Reading, Richard P., Tim W. Clark, John H. Seebeck, and Jennie Pearce. 1996. Habitat Suitability Index Model for the Eastern Barred Bandicoot, Perameles gimnii. Wildlife Research. 23:221-35. Saunders, D A, RJ. Hobbs, and C.R. Margules. 1991. Biological consequences of ecosystem fragmentation: a review. Conservation Biology 5:18-32. Slankis, V. 1973. Hormonal relationships in mycorrhizal development. Pages 231-298 in G. C. Marks and T. T. Kozlowski, eds. Ectomycorrhizae, their ecology and physiology. Academic Press, New York. Soule, M.E., and K.A. Kohm (eds.) 1989. Research priorities for conservation biology. Island Press, Washington, DC 97 pp. Stith, B.M., J.W. Fitzpatrick, G.E. Woolfenden, and B. Pranty. 1996. Classification and conservation of metapopulations. a case study of the Florida Scrub Jay. Pages 187215 in D R. McCullough. (ed) Metapopulations and Wildlife Conservation. Island Press, Washington. Tappe, Philip A., John W. Edwards, and David C. Guynn, Jr. 1993. Capture methodology and density estimates of southeastern fox squirrels (Sciurus niger). Pages 71-84 in N. D. Moncrief, J. W. Edwards, and P. A. Tappe, eds. Proceedings of the Second Symposium of Southeastern Fox Squirrels, Sciurus niger. Virginia Museum of Natural History Special Publication No. 1 . 84pp. Tappe, Philip A., and David C. Guynn. 1998. Southeastern fox squirrels: r-or K-selected implications for management. Pages 239-248. in Michael A. Steele, Joseph R. Merritt, and David A. Zegers, eds. Ecology and evolutionary biology of tree squirrels. Virginia Museum of Natural History, Special Publication Number 6. Teitge, Roberta M. 1992. Wildlife and golf courses. Pages 441-478 in J. C. Balogh and W. J. Walker, eds. Golf course management and construction. Environmental issues. Lewis Publishers, Boca Raton, FL. Terman, Max R. 1997. Natural links: naturalistic golf courses as wildlife habitat. Landscape and Urban Planning 38: 183-197.

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130 Thomasma, Linda E., Thomas D. Drummer, and Rolf 0. Peterson. 1991. Testing the habitat suitability index model for the fisher. Wildlife Society Bulletin. 19(3):291-297. Thompson, D. C. 1978. Regulation of a northern grey squirrel (Sciurus carolinensis) population. Ecology. 59: 708-715. Trappe, J. M, and C. Maser. 1977. Ectomycorrhizal fungi: interactions of mushrooms and truffles with beasts and trees. Pages 165-179 in T. Walters, ed. Mushrooms and man. USDA Forest Service, U. S. Government Printing Office, Washington, D. C. Turner, M. G. 1989. Landscape ecology: the effect of pattern on process. Annual Review of Ecology and Systematics. 20: 171-197. United States Fish and Wildlife Service. 1980. Flabitat evaluation procedures (HEP). United States Fish and Wildlife Service, Division of Ecological Services Manual 102. United States Fish and Wildlife Service. 1981. Standards for the development of habitat suitability index models. United States Fish and Wildlife Service, Division of Ecological Services Manual 103. Van Home, B. 1983. Density as a misleading indicator of habitat quality. Journal of Wildlife Management. 47(4): 893 -901. Weigl, P. D., M. A. Steele, L. J. Sherman, J. C. Ha, and T. S. Sharpe. 1989. The ecology of the fox squirrel in North Carolina: implications for survival in the Southeast. Tall Timbers Res. Stn. Publ. No. 24:1-93. Weston, John. 1990. Using native plants in the golf course landscape. USA Green Section Record. 28:12-15. Weston, John. 1994. Fire as a landscape management tool. USA Green Section Record. 32:14-16. White, G. C, and R. A. Garrott. 1990. Analysis of wildlife radio-tracking data. Academic Press, San Diego, Calif. 383pp. Williams, C. B. 1964. Patterns in the balance of nature. Academic Press, London. Williams, K. S., and S. R Humphrey.. 1979. Distribution and status of the endangered Big Cypress fox squirrel {Sciurus niger avicennia) in Florida. Fla. Sci. 42:201:205. Worton, B. J. 1989. Kernel methods for estimating the utilization distribution in homerange studies. Ecology 70:164-168.

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131 Worton, B. J. 1995. Using Monte Carlo simulation to evaluate kernel-based home range estimators. J. Wildl. Manage. 59:794-800. Wray, S., W. J. Cresswell, P.C.L. White, and S. Harris. 1992. What, if anything, is a core area? An analysis of the problems of describing internal range configurations. Pages 256-271 in Imants G. Priede and Susan M. Swift, eds. Wddlife telemetry: Remote monitoring and tracking of animals. Horwood. New York.

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BIOGRAPHICAL SKETCH Rebecca Selfridge Ditgen received a Bachelor of Arts degree in political science and education and a Master of Science in geography from the University of Kansas. For her master's thesis she examined the glacial history of the Kansan Age glacial deposits of the Hesper Plain in northeastern Kansas. She completed coursework and qualifiying examinations in the Ph.D. program in geography at the University of Wisconsin, Madison, in 1983. The emphasis of her studies was biogeography and the ecology of small mammals. She has worked for the National Park Service in Colorado and Florida. Rebecca began her Ph D program at the University of Florida in 1994, spending 2 .5 years studying fox squirrels in southwest Florida. 132

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I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is folly adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophy. Stephen R. Hun Professor of Wildlife Ecology and Conservation I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophy. iisenberg Katharme Ordway Pfofessdr of Ecosystem Conservation I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a dissertation for the degree of Doctor of Philoso^h^. /J Melvin Sunquist Associate Professor of Wildlife Ecology and Conservation I certify that I have read this study and that in my opinion it conforms to acceptable standards of scholarly presentation and is folly adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophv Mark Brown Assistant Professor of Environmental Engineering Sciences This dissertation was submitted to the Graduate Faculty of the College of Agriculture and to the Graduate School and was accented as partial fulfillment of the requirements for the degree of Doctor of Philosophy: J December, 1999 Dean, College of Agriculture Dean, Graduate School


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