Citation
Microhabitat use by Podomys floridanus in the high pine lands of Putnam County, Florida

Material Information

Title:
Microhabitat use by Podomys floridanus in the high pine lands of Putnam County, Florida
Creator:
Jones, Cheri A., 1957-
Publication Date:
Language:
English
Physical Description:
xii, 160 leaves : ill. ; 29 cm.

Subjects

Subjects / Keywords:
Mice -- Florida ( lcsh )
Putnam County ( local )
Tortoises ( jstor )
Burrows ( jstor )
Mice ( jstor )
Genre:
bibliography ( marcgt )
theses ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis (Ph. D.)--University of Florida, 1990.
Bibliography:
Includes bibliographical references (leaves 146-158).
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by Cheri A. Jones.

Record Information

Source Institution:
University of Florida
Holding Location:
University of Florida
Rights Management:
Copyright [name of dissertation author]. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
Resource Identifier:
022465149 ( ALEPH )
AHH1974 ( NOTIS )
79166853 ( OCLC )

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Full Text









MICROHABITAT USE BY PODOMYS FLORIDANUS IN THE HIGH
PINE LANDS OF PUTNAM COUNTY, FLORIDA










By

CHERI A. JONES


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


1990






















Copyright 1990

by

Cheri Ann Jones













ACKNOWLEDGMENTS

I thank my chairman, J. F. Eisenberg, for providing

encouragement, advice, and equipment during this project.

Members of my committee (R. Franz, J. H. Kaufmann, R. A.

Kiltie, F. E. Putz, and J. G. Robinson) made useful comments

and suggestions regarding this paper. John Eisenberg and

Dick Franz bear responsibility for introducing me to the

Ordway Preserve, where I formed a strong attachment to

Florida mice, gophers, and sandhills. I also benefited from

discussions with C. A. Gates, J. I. Glendinning, J. N.

Layne, and members of the Eco-lunch seminar group in the

Department of Zoology.

I am grateful to the Florida Game and Fresh Water Fish

Commission for issuing permits allowing me to trap Florida

mice and to establish a captive colony. I borrowed aquaria,

traps, and other equipment from J. F. Anderson, L. S. Fink,

W. H. Kern, R. F. Labisky, C. A. Langtimm, F. Nordlie, P.

Ryschkewitsch, and C. A. Woods. The Florida Foundation

provided funds for additional traps, flags, and other field

supplies. I thank T. Perry, the preserve caretaker, for

providing information about the history of Ordway and for

pulling my truck out of the sand more frequently than I care

to remember. Help in caring for captive animals, counting


iii








plants, and other forms of assistance were provided by R. A.

Drury, J. Dudley, J. F. Eisenberg, K. Eisenberg, J. Ellis,

N. Etheridge, R. Franz, S. E. Franz, B. Furlow, A. Gannon,

M. K. Lacy, C. A. Langtimm, A. Mahler, M. Marquez, R. Mor-

rison, C. Partin, L. Partin, J. R. Thomson, and W. W. Tim-

merman. I thank R. Franz and D. W. Hall, who patiently

identified plant specimens and taught me much about the high

pine flora. J. Boucher, J. I. Glendinning and K. M. Portier

helped solve statistical and computer problems.

Burns on the Ordway-Swisher Preserve were conducted by

J. F. Eisenberg, R. Franz, T. Perry, and M. E. Sunquist.

These burns would be impossible without additional help; I

thank all those volunteers (including R. Burke, P. Carnell,

B. Charest, C. K. Dodd, Jr., R. A. Drury, S. E. Franz, C. A.

Gates, H. Gold, L. Hay-Smith, C. A. Langtimm, M. E. Ludlow,

R. Morrison, D. Pearson, F. E. Putz, G. W. Tanner, K. K.

Timmerman, D. D. Wright, and students from the departments

of Botany, Wildlife, and Zoology) who cheerfully incinerated

my study sites and other sandhills.

Finally, special thanks are due to my parents, Clyde

and Charlene Jones, and my grandparents, Ellis and Maye

Owen, who have provided constant encouragement and moral

support throughout my academic career. I also am grateful

to R. A. Drury, P. Foster-Turley, R. and S. E. Franz, and C.

A. Langtimm, who offered friendship and support during my

years in Gainesville.

















TABLE OF CONTENTS


ACKNOWLEDGMENTS .

LIST OF TABLES .

LIST OF FIGURES .

ABSTRACT .

INTRODUCTION .

TAXONOMY AND SYSTEMATICS .

DISTRIBUTION .

MATERIALS AND METHODS .

USE OF TORTOISE BURROWS .

Dependence on Burrows .
Description of Burrows .
Podomys and the Burrow Community
Life History .

HOME RANGE .

Fluorescent Powder .
Trapping on Grids .
Trapping at Burrows .
Conclusions .

DIET .

Field Observations .
Laboratory Observations .
Acorn Preference Tests .. ..
Supplemental Feeding Experiment

ANALYSES OF VEGETATION .

Sampling Technique .
Results and Discussion .


......... iii
. vii
. Vii

. ix

. xi

. 1

. 4

. 7

. 10

. 18

. 18
. 26
. 31
. 34

. 40

. 41
. 44
. 51
. 65

. 68

. 68
. 70
. 72
. 78

. 84

. 84
. 89







PRESCRIBED BURNS . 100

Materials and Methods 100
Description of Procedure for Prescribed Burns 102
Results and Discussion 103
Conclusions . 129

SUMMARY AND DISCUSSION . 132

APPENDIX LIST OF PLANT SPECIES IDENTIFIED IN
VEGETATION SAMPLES ... 140

LITERATURE CITED . .. 146

BIOGRAPHICAL SKETCH ... .159













LIST OF TABLES


Table Page

1 Numbers of captures of non-target species in
traps at burrows (22,085 trapnights) and on
grids (11,300 trapnights) 24

2 Data describing distribution of burrows on SL
(top) and AC (bottom) with the probabilities
expected from the Poisson distribution (with
means of 1.5 and 1.7 burrows per square) .. 30

3 Longevity records exceeding 360 days for
Podomys floridanus at SL ... .39

4 Activity data from ten Podomys dusted with
fluorescent powder at SL ... 43

5 Home range data for 20 Podomys trapped on two
grids .... . 48

6 Capture data for 36 Podomys at Anderson-Cue 49

7 Capture data for the 62 adults whose home
ranges are shown on Fig. 9 .. 61

8 Persistence data for animals on grids 83

9 Habitat variables measured in each quadrat 86

10 Frequency (% = percent of quadrats in which
a species occurred) and density (# = average
number of stems per 314 m2 quadrat) of woody
species identified at SL ... .90

11 Frequency (% = percent of quadrats in which
a species occurred) and density (# = average
number of stems per 314 m2 quadrat) of woody
plant species identified at BP in 1988 .. .92

12 Habitat profile of Podomys at all BP and SL
burrows where vegetation was sampled .. 94


vii








13 Number (%) of vegetation samples reassigned
according to discriminant function analysis
to two categories of total number of captures
(ALL) of Podomys .. 96

14 Burrows on burned sandhills ..... 111

15 Burrows on unburned sandhills ......... 112

16 Comparison of capture indices for Podomys
at burrows on burned and unburned sandhills 113

17 Comparison of capture indices for Podomys at
burrows on 3 burned sandhills 115

18 Survival of Podomys following three prescribed
burns on the Ordway Preserve .120

19 Comparison of trapping effort on unburned and
burned sandhills, where TN is the total number
of trapnights and trapping effort is expressed
as trapnight/capture/ha (TN/CAP/HA) ... 122

20 Residence of young Podomys on the SL sandhill,
where N is the total number of young marked
and R is the percentage of marked young that
became residents on that sandhill ....... .123

21 Responses of Peromyscus species to fire .... 125


viii














LIST OF FIGURES

Figure Page

1 Distribution of Podomys floridanus 8

2 Distribution of Podomys on the Ordway/Swisher
Preserve . ... .. .14

3 Trapping success (%) and minimum numbers of
individuals (MNI) (adults and subadults) of
Podomys at burrows on the Smith Lake Sandhill 22

4 Trapping success (expressed as the total num-
ber of captures/adjusted trapnights) for Podo-
mvs on the AC (top) and SL (bottom) grids .. 27

5 Numbers of juveniles (dots) and subadults
(circles) marked on the SL sandhill, expressed
as minimum numbers of individuals/number of
trapnights (top) and as numbers of individuals
(bottom) . ... .. 36

6 Reproductive data for Podomys trapped on the
SL sandhill, 1983-1988: the proportion of
adult females who were pregnant (top) and the
proportion of males who were scrotal (bottom)
are shown .... .37

7 Trapping data for Podomys on the ACI, ACII,
and SL grids . ... 45

8 Home ranges of Podomys trapped on the ACI grid
in May 1987 . 52

9 Annual home ranges of 62 Podomys on the SL sand-
hill, 1983-1988. ... .55

10 Minimum numbers of individuals (MNI) on the ACI,
ACII, and SL grids .. 81

11 Apparent densities (expressed as minimum numbers
of individuals/ha) of Podomys on the burned and
unburned areas of the Blue Pond sandhill ..... 105








12 Apparent densities (expressed as minimum numbers
of individuals/ha) of Podomys on the burned and
unburned areas of the Longleaf Pine sandhill 107

13 Apparent densities (expressed as minimum numbers
of individual/ha) of Podomys on the burned and
unburned areas of the Smith Lake Sandhill 109

14 Home ranges of eight Podomys on the SL sandhill
before (stippled) and after (hatched) the 1985
fire . 117













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

MICROHABITAT USE BY PODOMYS FLORIDANUS IN THE HIGH
PINE LANDS OF PUTNAM COUNTY, FLORIDA

By

Cheri A. Jones

May 1990

Chairman: John F. Eisenberg
Major Department: Zoology

The purpose of this project was to examine the use of

tortoise burrows (Gopherus polyphemus) by the Florida mouse

(Podomys floridanus) in high pine (longleaf pine sandhills)

on the Katharine Ordway Preserve/Swisher Memorial Sanctuary

in Putnam County, Florida. Mark-and-recapture techniques

were used at tortoise burrows and on grids (10000 m2) in

order to describe burrow use by mice and to measure home

ranges. I also performed a food supplementation experiment

to test the hypothesis that population sizes were food-

limited. Finally, I examined how local distributions of

Florida mice were correlated with vegetation structure and

affected by prescribed burns.

Podomys floridanus are commensals of Gopherus in high

pine habitats at Ordway, as indicated by trapping success,

escape responses, burrow excavations, and observations of








mice in the field. In five years of trapping at burrows on

a single sandhill, 255 mice were marked. Of this total, 32%

were captured only once and 8.6% survived for 360-920 days.

Reproductive activity was concentrated in August, September,

and October. Florida mice appeared to be opportunistic

feeders, accepting a wide variety of fruits, seeds, and

insects. Populations appeared to be limited by factors

other than food. Mean annual home ranges were 2601 m2 for

adult females, whose ranges were mutually exclusive. Home

ranges of adult males were significantly larger (4042 m2)

and overlapped with those of other adults. Numbers of

captures at burrows were best correlated with large numbers

of herbaceous species and high stem counts of Pinus palus-

tris, Quercus laevis, and Diospyros virginiana; there was

little correlation with ground cover characteristics.

Immediate responses of Podomys to prescribed burns were not

spectacular, but in the long term, populations appeared to

be more stable on sites that were burned periodically than

on areas not burned. Implications for conservation of

Podomys on the Ordway Preserve also were discussed.


xii













INTRODUCTION


An understanding of home range, diet, burrow use, and

other interactions between an animal and its physical en-

vironment is crucial for the management and conservation of

threatened and endangered species. However, little is known

about the ecology of Florida's only endemic mammal, the

Florida or gopher mouse. This mouse, Podomys floridanus

(Rodentia: Muridae), is listed as Threatened by the Florida

Committee on Rare and Endangered Plants and Animals (Layne,

1978) and as a Species of Special Concern by the Florida

Game and Fresh Water Fish Commission (1987). Podomys flori-

danus lives in a narrow range of habitats; my study, and

those by Layne at the Archbold Biological Station in High-

lands County, suggest that P. floridanus populations are

higher in the southern sand pine scrub than in the high pine

lands (also known as turkey oak-longleaf pine sandhills or

high pine hills). The special listing of P. floridanus is

due to destruction of high pine and scrub habitats and to

the decline of gopher tortoise (Gopherus polyphemus) popula-

tions. Nash (1895) stated that high pine was the most

extensive of the five habitats he recognized in Florida; by










1982 approximately 57% of this habitat had been lost (Auf-

fenberg and Franz, 1982).

In the high pine lands of northern Florida, the Florida

mouse is associated closely with burrows of the gopher

tortoise (Blair and Kilby, 1936; Humphrey et al., 1985;

Brand, 1987). My preliminary field studies showed that

populations of Florida mice were relatively high and stable

on certain sandhills, whereas populations fluctuated to a

greater degree on other sites. I was intrigued by the

differences I observed in these populations. How did these

sandhills differ from each other? Why were certain tortoise

burrows favored nesting sites, year after year, whereas

other burrows never were used? In an attempt to determine

microhabitat preferences in the high pine habitats, I tried

five approaches:


1) Do Podomys in the high pine depend exclusively on
tortoise burrows? Mouse-use of tortoise burrows
was monitored over a five-year period so that
long-term "burrow histories" could be constructed.


2) How large are home ranges in the high pine? Home
range areas of Podomys in high pine were measured.

3) Do dietary data explain why some tortoise burrows
are preferred over others? Diets of these mice
were observed in the field and laboratory. A
supplemental feeding experiment was performed to
test the hypothesis that population size is food-
limited (i.e., does increased food supply produce
an increase in numbers and body size of mice and
an increase in the length of the breeding sea-
son?).








3

4) Is the local distribution of these mice predic-
table? Physical parameters of burrows, and the
vegetation surrounding them, were quantified.

5) Do populations respond positively to burns?
Effects of prescribed burns on populations of
Podomys were determined.













TAXONOMY AND SYSTEMATICS


Podomys floridanus is the only mammalian species en-

demic to Florida. The species was described as Hesperomys

floridanus by Frank Chapman in 1889, with the type locality

in Gainesville, Alachua County.

Podomys floridanus has long been recognized as distinct

from North American species of Peromyscus. Osgood (1909)

considered Podomys a distinct subgenus within the genus

Peromyscus. Apparently Hooper (1958) was the first to

suggest that the closest relatives of Podomys were not the

cotton mouse and other local Peromyscus, but peromyscines in

Central America. In Carleton's (1980) major revision of the

genus Peromyscus, Podomys was raised from subgenus to genus.

Carleton suggested that Podomys shared a common ancestor

with Habromys and Neotomodon from Mexico and Guatemala;

these three genera grouped together consistently in his

cladistic analysis of 79 morphological characters.

The proposed affinity between Podomys and mice from

Central America is supported by three lines of evidence:

1) Reproductive tract. The baculum of Podomys is
relatively small and slender, compared to that of
northern Peromyscus. Linzey and Layne (1969)
examined additional features of the male reproduc-
tive tract and noted reduced vesicular and










prostate glands, enlarged ampullary glands, and
other features that suggested affinity with Habro-
mys.

2) Ectoparasites. The "typical and only common flea"
of Podomys, described by Johnson and Layne (1961),
is Polygenis floridanus, which most closely resem-
bles a species found in South America. According
to Wenzel and Tipton (1966: 718), "Polygenis is an
"expanding" South American genus which has, quite
clearly, dispersed into Middle America along with
complex-penis-type Cricetinae and caviomorph
rodents from South America".

3) Electrophoresis. In their analysis of genic
variation among Peromyscus, Smith et al. (1973)
considered Podomys one of the least variable
species of the group. One possible explanation
for this low variability is the maintenance of low
numbers in a relictual population.

The fossil record of the Florida mouse is limited to

six Rancholabrean assemblages in this state, so there is not

much light shed on the distribution of Podomys prior to the

Illinoisan. However, a western affinity might be indicated

by a Pleistocene species (Peromyscus oklahomensis) from the

Illinoisan and Sangamonian of Oklahoma and Texas, that might

be related to Podomys (Stephens, 1960). Unfortunately, only

a few isolated teeth of this extinct species have been

found.

Podomys is not unique among the flora and fauna of

Florida in its apparent affinity to Central America. Several

plants (such as Opuntia, the prickly pear) and several

reptiles, birds, and mammals are considered species of

western affinity. Some, such as Gopherus polyphemus, show a

disjunct distribution (Blair, 1958). Among the mammals, Webb

and Wilkins (1984) found that 15-28% of the mammals from








6

five Pleistocene sites in Florida were neotropical species.

They suggested that the arrival of most of these species

could be explained by a Gulf Coastal Corridor, which served

as a pathway for southern and western immigrants in the

early Pleistocene.













DISTRIBUTION


In recent times P. floridanus occurred from St. Johns,

Clay, Alachua, Putnam, Suwannee, and Taylor counties south

to Sarasota, Highlands, and Dade counties, with isolated

populations in Franklin County and on Merritt Island (Os-

good, 1909; Layne, 1978; Stout et al., 1979). A survey of

the present distribution of Podomys is urgently needed.

Podomys is the characteristic mouse of some of the most

xeric habitats in Florida. It is most common in sand pine

scrub and high pine, although it also is found in slash

pine-turkey oak and coastal strand associations (Bangs,

1898; Layne, 1978; U.S. Dept. Agric., 1980). On Fig. 1, I

have mapped the distribution of scrub and high pine, as

indicated by the Soil Conservation Service (U.S. Dept.

Agric., 1980), and localities from which Podomvs have been

reported (Layne, 1978). Clearly there is a close associa-

tion between Podomys and these two habitats. Layne (in

press) has suggested that the scrub was the ancestral home

of this mouse, with the high pine a secondary habitat.

In some areas, P. floridanus and P. gossypinus appear

to coexist (Layne, 1970). In Putnam County, however, trapp-

ing by Suzanne Brand (1987), John Eisenberg (1983; 1988),






































80 km


as 0


Fig. 1. Distribution of Podomys floridanus (triangle
designates type locality). Stippling represents
distribution of high pine and scrub habitats.








9

Eric Milstrey (1987), and me has indicated a clear separa-

tion between these two species, with P. aossypinus occurring

only in lower, wetter microhabitats, and Podomys only in

sandhills, or along edges between sandhills and old pastures

or sandhills and xeric hammocks.













MATERIALS AND METHODS


I conducted this research on the Katharine Ordway

Preserve-Carl S. Swisher Memorial Sanctuary, located about

4 km E of Melrose in Putnam County. The preserve, 37 km2 in

size, is administered jointly by the Florida Museum of Na-

tural History, the University of Florida's School of Forest

Resources and Conservation, and the Nature Conservancy.

Approximately one-third of the Ordway Preserve is

covered by high pine land (sensu Nash, 1895). This habitat

is characterized by an open canopy of longleaf pine (Pinus

palustris) and turkey oak (Quercus laevis), although other

trees, including sand pine (P. clausa), Chapman's oak (Q.

chapmanii), and bluejack (Q. incana), also are present. The

understory consists of scattered shrubs, and herbs and

grasses. On Ordway the characteristic grass of the high

pine land is wiregrass (Aristida stricta); other common

grasses include pineywoods dropseed (Sporobolus junceus),

six species of bluestem (Andropogon spp.), and three species

of threeawn (Aristida spp.). Additional grasses identified

on the preserve are included in the list compiled by David

W. Hall and Richard Franz (unpubl. data). Reindeer moss









(Cladonia and Cladina spp.) is common. The herbs I

identified on my study sites are listed in the Appendix.

The soils of high pine are sandy and well-drained.

There are many patches of bare ground, which become covered

with oak leaves, pine needles, tree branches, and other

debris in the absence of fire. Under the pines, the needle

litter is several inches deep; in areas where the turkey

oaks predominate, leaf litter is distributed unevenly by the

wind. Burrows of the gopher tortoise (Gopherus polyphemus)

are prominent features on this landscape, although popula-

tion densities probably were higher in coastal areas before

human development (Carr, 1940; Auffenberg and Franz, 1982;

Cox et al., 1987).

Other mammals found on the high pine land are the

opossum (Didelphis virainianus), pocket gopher (Geomys

pinetis), fox squirrel (Sciurus niger), flying squirrel

(Glaucomys volans), armadillo (Dasypus novemcinctus), rac-

coon (Procyon lotor), and white-tailed deer (Odocoileus

virainianus). Eisenberg (1988) and Franz (1989) compiled

lists of mammals known from the various habitats of Ordway.

Carr (1940) listed the amphibians and reptiles of high pine

habitats. On my study sites, I saw gopher frogs (Rana

areolata), oak toads (Bufo quercicus), fence lizards (Scelo-

porus undulatus), six-lined racerunners (Cnemidophorus

sexlineatus), pine snakes (Pituophis melanoleucus), crowned

snakes (Tantilla spp.), southern black racers (Coluber









constrictor), coachwhips (Masticophis flagellum), coral

snakes (Micrurus fulvius), pigmy rattlesnakes (Sistrurus

miliarius), eastern diamondbacks (Crotalus adamanteus), and

tortoises (G. polyphemus). Birds common to high pine were

listed by Nelson (1952), who thought the avifauna of the

high pine did not differ substantially from those of oak

hammocks and other nearby communities. However, Maehr et

al. (1982) suggested that the high pine might support a

greater density of birds than flatwoods and xeric hammocks.

I did not attempt to identify birds on my study sites in

detail.

On the Ordway Preserve, all locations at which P.

floridanus have been found are in high pine habitats, or at

the ecotone between high pine and hammock or high pine and

old pasture (Brand, 1987; Eisenberg, 1988) (Fig. 2). I

trapped mice on four independent high-pine sites, designated

Anderson-Cue (AC), Blue Pond (BP), Longleaf Pine Pasture

(LP), and Smith Lake (SL). The history of fire on these

sites is described below in the Prescribed Burns section.

Smith Lake is my longterm study site; Eisenberg and Franz

started research projects there in 1983, marking tortoise

burrows and trapping Florida mice, gopher frogs, and other

burrow inhabitants. I trapped mice at SL from 1984 through

1988. SL is approximately 16.8 ha in size. Vegetation

consists of P. palustris, Q. laevis, Q. chapmanii, Q. qemin-

ata, and grasses and forbs characteristic of the high pine.













4-4 wQ)


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15

The Anderson-Cue sandhill is the highest in elevation of the

four sites; typical vegetation there includes P. palustris

and Q. laevis, with fewer Q. geminata than on SL. I worked

there from 1986 to 1988. AC was burned in the winters of

1983, 1986, and 1989. It was not included in my study of

prescribed burns because I did no pre-burn trapping there.

The Longleaf Pine Pasture Sandhill is south of the pine

pasture surrounding the old Swisher Estate (Fig. 2). Part

of this sandhill, the most disturbed of my study areas, has

been planted in slash pine. Dominant plants here include

maidencane (Panicum hemitomon) and centipede grass (Eremoch-

loa ophiuroides), which are not typical of natural high pine

vegetation. There are many turkey oaks and several large

patches of gopher apple, but fewer longleaf pines, pawpaws,

and other high pine species. I worked on LP for only one

trapping season for the purpose of gathering additional

information regarding prescribed burns. The fourth study

site, Blue Pond, was logged intensively. There are few

longleaf pines, and turkey oaks are by far the most numerous

trees. Ground cover is sparse in some areas but rather

dense in others. Species composition of the ground cover

appeared similar to that of SL.

As described in the following chapters, I trapped E.

floridanus at tortoise burrows and on grids using standard

mark-and-recapture techniques (Davis, 1956). While trapping

at burrows, I numbered, marked, and mapped each burrow, and








16
then placed a pair of Sherman traps (baited with oatmeal) at

the entrance, or just inside the mouth, of the burrow.

Animals were marked by toe-clipping and then weighed. I

also noted sex, reproductive condition, and presence of

ectoparasites. Animals were released near the burrow en-

trance, and in most cases I recorded whether they entered

the burrow or ran elsewhere. I collected the same data on

grids. I placed one grid (designated SL) on my study site

on the south side of SL; two more grids (ACI and ACII) were

set north and east of the Anderson-Cue ponds. Grids were

more than 100 m apart to reduce movement of animals between

grids. Each grid consisted of 10 columns and 10 rows 10 m

apart (area = 10000 m2), with a single Sherman at each

intersection. Each tortoise burrow and grid location was

given a unique number and marked with forestry flags. On

the burrow flags I also attached an aluminum tag (Al Tag,

Forestry Suppliers, 205 W. Rankin Street, Jackson, Missis-

sippi 39204) bearing the burrow number. These tags survived

all but the hottest burns, thus serving as more permanent

markers of burrow locations.

Traps were opened near sundown and checked at or short-

ly following sunrise. Because of temperature extremes,

morning condensation, and (in some months) heavy rain, I

found it necessary to check traps in the morning as quickly

as possible. Animals developed hypothermia when ambient

temperatures reached 10 C; I extended the trapping season










into November by insulating traps with a handful of excel-

sior, wood shavings used in packing glassware. Mice com-

press the excelsior into a pad on top of the treadle.

Spanish moss also can be used, but I found the wood to be

superior because it does not absorb moisture. I did not

trap in December or January; fieldwork resumed when warm

weather resumed (February-April).

I measured trapping effort as trapnights (one trapnight

being one trap/night). Results of the vegetation analyses

were run on SYSTAT at the IFAS Consulting Center at the

University of Florida; other statistics were performed on a

hand-held calculator. Statistical methods are described in

each chapter.













USE OF TORTOISE BURROWS


Dependence on Burrows

Previous students of peromyscine behavior and morphol-

ogy concluded that Podomys was primarily a terrestrial

species; although capable of climbing and digging, Podomys

climbed and dug less than several species of Peromyscus to

which it was compared (King, 1968; Layne 1970; Layne and

Ehrhart, 1970; Siegel and Van Meter, 1973). In some areas,

these mice seemed capable of excavating their own burrows;

Lee (1968) stated that P. floridanus built burrows greater

than 120 cm in depth in scrub habitats, and Starner (1956)

trapped one animal in a small burrow that she thought might

have been dug by the mouse (in the ecotone between the mesic

hammock and longleaf pine flatwoods at San Felasco Hammock,

Alachua County). Layne and Ehrhart (1970) suggested that

Podomys rarely dug its own burrows, but Layne (in press)

noted high populations of Florida mice in scrubs where

tortoises were absent or occurred in low numbers. However,

observations by Eisenberg (1983), Brand (1987), and myself

suggested a much closer relationship between Podomys and

Gopherus in the high pine in Putnam County.










Blair and Kilby (1936) were the first to note in the

literature that Podomys floridanus used burrows of Gopherus

polyphemus in northern and central Florida. They saw mice

using active burrows (i.e., burrows that had tortoises in

them) in an old field in Alachua County. Three of the 15

mice captured by Starner (1956) were taken from traps placed

inside entrances of tortoise burrows. Milstrey (1987:115)

reported that "Florida mice were continuously present" in

many tortoise burrows at his study site in Putnam County.

Johnson and Layne (1961) also noted that Podomys inhabited

tortoise burrows, and possibly burrows of pocket gophers

(Geomys pinetis). At other sites Podomys used burrows

constructed by Peromyscus polionotus, Sigmodon hispidus, G.

pinetis, and Dasypus novemcinctus (Gentry and Smith, 1968;

Layne, in press). However, in my 33,385 trapnights on

burrows and grids at Ordway I have observed Florida mice

entering small holes of unknown origin only three times

(twice on SL and once on AC), and I have no evidence that

they inhabited logs or other shelters.

I found that mice almost always ran down tortoise

burrows when released. For example, I tallied escape re-

sponses of 18 Podomys caught on the AC grid (and therefore

released at grid coordinates, not at burrows). Out of 35

captures, mice ran down tortoise burrows 25 times; they hid

in the base of turkey oaks or in fallen logs and other

debris 10 times. In my regular trapping sessions, I never








20
saw mice climb vegetation as an escape response; Eisenberg

(pers. comm.) noted one such response in 1989. However, my

field observations have confirmed climbing ability by these

mice (Jones, 1989, and Home Range below). In T-maze tests

of arboreality performed by C. A. Langtimm at SL (in which a

T-maze was propped against an oak), most mice jumped off the

T, but the animals who chose to climb oaks reached heights

of 8 m (C. A. Langtimm, unpubl. data). Clearly these ani-

mals are capable of climbing, even though the time spent on

this activity in any one night remains unknown.

In preliminary studies on the Ordway Preserve, Eisen-

berg (1983) reported that trapping success was much higher

if traps were placed at the mouths of tortoise burrows (33%)

than on transects (0.4%). Humphrey et al. (1985) caught 113

mice in 1,144 trapnights at 51 burrows, also at Ordway. In

146 trapnights on transects--in which traps were not placed

at burrows--I caught no mice. At the SL burrows in August,

September, October, and November 1983, Eisenberg (unpubl.

data) had trapping successes of 35, 43, 62, and 44%, respec-

tively. I calculated trapping success using adjusted trap-

nights, which indicated how many traps were available after

subtracting half the number of traps sprung and containing

non-target species from the total number of traps set. My

trap success for Podomys ranged from 0 to 16%, with SL being

the most successful (2-16% with a mean of 7%) (Fig. 3).

Variations in total trapnights (within and among sites)















W)

'-I








4J

00





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reflect my attempt to maximize sample size by trapping at

all tortoise burrows on each sandhill until burrows disap-

peared.

On all four sandhills, traps at burrows occasionally

were sprung by falling branches or by insects removing bait.

A large number of sprung traps was an occasional problem at

AC, but a more major one at BP, particularly on the north

side. There I identified tracks of armadillos, raccoons,

and dogs, but I suspected dogs were the major culprits. One

morning I found two traps covered with urine and several

times I saw dogs on the sandhill, which was the closest of

my sites to the periphery of the preserve. It is also worth

noting that I found more carcasses of turtles and tortoises

here than on any other study site. Following a suggestion

of M. E. Sunquist, I fashioned "croquet hoops" out of alu-

minum welding rods and placed them around the traps. They

helped decrease the number of traps moved, but did not

always prevent closing of trap doors.

In traps at burrows, the most frequently-captured non-

target species was Rana areolata (Table 1). At SL, from

September 1984 to the end of the study, gopher frogs were

captured 105 times. Other occasional captures were camel

crickets (Ceuthophilus sp.), centipedes, southern toads

(Bufo terrestris, once on AC, LP, and SL), six-lined race-

runners (Cnemidophorus sexlineatus), and a young rabbit

(presumably Svlvilaqus floridanus, on SL). All of these








24

TABLE 1. Numbers of captures of non-target species in traps
at burrows (22,085 trapnights) and on grids (11,300 trap-
nights). Captures of insects and centipedes were not
counted. Scientific names are listed in text.


Burrows

camel crickets

1 spider

3 southern toad

129 gopher frog



3 six-lined racerunner

1 cotton mouse

38 flying squirrel

1 rabbit


Grids

grasshoppers

cockroaches

centipedes

3 six-lined

racerunner

2 fence lizard

1 black racer

2 rice rat

3 cotton rat

148 flying squirrel










species are known to use Gopherus burrows (Jackson and

Milstrey, 1989). In 1988 I began catching flying squirrels

(Glaucomys volans) at the AC and SL burrows; at SL I cap-

tured squirrels 27 times from May to October, 15 times in

May alone. These results must indicate some change in the

Glaucomys population that resulted in additional terrestrial

foraging, or perhaps they reflect learned behavior of ani-

mals who found oats at traps on grids. At any rate, I

observed only one flying squirrel enter a burrow upon re-

lease, and I do not believe they reside in tortoise burrows

on these sandhills.

At study sites other than Ordway, additional murids,

including P. gossypinus and P. polionotus, visit tortoise

burrows (Jackson and Milstrey, 1989). In 22,085 trapnights

at burrows on the AC,BP, LP, and SL sandhills I caught only

one cotton mouse (P. gossypinus, on AC). In other Ordway

studies, Eisenberg (1983) and Brand (1987) found Podomys in

the high pine and Peromyscus in the hammocks and flatwoods.

Clearly, Podomys is the only rodent that regularly inhabits

tortoise burrows in these sandhills. At burrows in other

Ordway habitats, the absence or lower numbers of Podomys

might represent a case of competitive exclusion; after the

1987 drought Eisenberg (pers. comm.) reported trapping

Florida mice in hammocks closer to lakes than in previous

years, and Peromyscus were absent.







26

For comparative purposes, trapping results on two grids

on AC and one grid on SL also are shown (Fig. 4). Trapping

success on grids ranged from 0 to 7% (AC) and from 0 to 6%

(SL); mean trapping successes were 0.47%, 0.70%, and 3.6%

for ACI, ACII, and SL. Traps on grids were more likely to

be sprung by rain, falling branches, pine cones, and rac-

coons than were traps at burrows. Only one Florida mouse

(female 18, a juvenile known to be alive 36 days at SL) was

found on a grid and not during burrow trapping. Flying

squirrels were caught more frequently (Table 1), usually at

traps near the bases of pines, oaks, or dead trees. One

cotton rat (Siqmodon hispidus) was captured once at SL; this

was a juvenile male (which I marked) presumably dispersing

across the high pine. The other cotton rats and the rice

rats (Orvzomys palustris) were caught on the AC grids on 28

and 29 June 1988; their appearance on the grids might re-

flect drought-related dispersal out of the vegetation sur-

rounding the AC ponds. Centipedes occasionally were caught

on all grids. On ACI I also captured grasshoppers, cock-

roaches, fence lizards (Sceloporus undulatus), six-lined

racerunners (C. sexlineatus), and a black racer (Coluber

constrictor).



Description of Burrows

The size and configuration of tortoise burrows have

been described by Hallinan (1923) and Hansen (1964). Blair












---0-- ACI
-- ACII


I
%
0, ,S

M J J AS O M A M J J AS O N


1987


1988


MONTH


A M J J ASOM A M J J A S O N
1987 MONTH 1988
MONTH


Fig. 4. Trapping success (expressed as the total number of
captures/adjusted trapnights) for Podomys on the
AC (top) and SL (bottom) grids.







28

and Kilby (1936) noted the use of a small hole approximately

182 cm from the entrance of a burrow. However, the only

description of structures used by Podomys was that of Layne

(in press), who described nests, nest chambers, and small

tunnels found in two tortoise burrows in Alachua County.

Richard Franz and I excavated five burrows on Ordway in

order to further describe structures utilized by mice.

Typically, mice used small U-shaped tunnels constructed off

one side of the tortoise burrow, with both entrances to the

U opening into the main burrow. We also noted use of tun-

nels we dubbed "chimneys", which ran from the ceiling or

side of the burrow to an outside opening a meter or two past

the main entrance. In all five burrows, mouse activity

appeared concentrated in the upper 2 m of the burrow.

Additional details of the actual excavations were discussed

by Jones and Franz (in prep.). Chimneys allowed occupation

of burrows after the main entrance has collapsed, and might

serve as escape routes from predators. Although not all

burrows have chimneys (at my four study sites from 6.8% (AC)

to 26.9% (BP) of the burrows had chimneys), the presence of

chimneys is an indicator of past or present occupation of

tortoise burrows by Podomys. Exploratory trapping for mice

in the sandhills should not omit observation of these small

holes.

To determine the pattern of dispersion of tortoise

burrows on the SL sandhill, I used methods described by







29

Brower and Zar (1984). In this procedure, a grid is placed

over a map of the study area and numbers of squares contain-

ing no burrows, squares containing one burrow, and etc., are

counted. I used a grid with squares 35 mm by 35 mm (which

corresponds to 1225 m2 on the map). My SL study site con-

sisted of 93 squares with a mean of 1.46 burrows per square

(or 0.11 burrow/m2). I included all 136 burrows monitored

from 1983 through 1988, without consideration of burrow

classification or tortoise demography; that is, I considered

only burrow locations without burrow size, duration of

residence, or other characteristics. The data fit the

Poisson distribution, leading to the conclusion that the 136

burrows on SL are randomly distributed (Table 2). Another

method to determine whether distribution is random is the X2

test, for which I obtained a value of 0.547 (not signifi-

cantly different from random at p = 0.05). As a third me-

thod, I tried the Morisita's index of dispersion, which is

calculated as I = n [(EX2-N)/(N(N-1))] where n is the number

of squares, N is the total number of burrows counted on n

squares, and ZX2 is the squares of the numbers of burrows

per plots for all plots. A value of 1.0 for I indicates

random dispersion; value of 0 indicates uniform dispersion.

I obtained I = 0.95, which indicates random dispersion.

Finally, I calculated the ratio of observed to expected

density; I obtained a value of 0.94, which is close to the

value of 1.00 which again indicates random distribution









TABLE 2. Data describing distribution of burrows on SL
(top) and AC (bottom) with the probabilities expected from
the Poisson distribution (with means of 1.5 and 1.7 burrows
per square). X indicates number of burrows per square; f(X)
indicates the observed frequency; p(X) is the observed
probability; and P(X) the expected Poisson probability.


X f(XIl pX_ P(X)

0 20 0.215 0.223

1 33 0.354 0.335

2 24 0.258 0.251

3 10 0.108 0.126

4 5 0.054 0.047

5 1 0.011 0.014


0 15 0.259 0.183

1 13 0.224 0.311

2 13 0.224 0.264

3 12 0.207 0.150

4 3 0.052 0.064

5 0 0 0.022

6 0 0 0.006


2 0.035


0.001









(Brower and Zar, 1984). The 101 AC burrows also were dis-

persed randomly (X2 = 5, significant at p = 0.10; I = 0.98;

ratio of observed to expected = 1.29).

Observations of dispersion of other gopher tortoise

burrows vary. McRae et al. (1981) and Auffenberg and Franz

(1982) recognized burrows clumped as colonies, whereas

others reported random distributions (e.g., Kushlan, 1982;

Kushlan and Mazzotti, 1984). Whether these discrepancies

reflect variations in tortoise behavior or in data collec-

tion remains unknown.



Podomys and the Burrow Community

In the high pine, Gopherus burrows provide shelter from

the fires and extreme fluctuations in temperature and rain-

fall that characterize this environment. Burrows of the

southeastern pocket gopher (Geomys pinetis) also are common

in the high pine, but the two types of burrows are used by

different species (Funderburg and Lee, 1968). Eisenberg

(1983) suggested that G. polyphemus could be considered a

keystone species in the high pine due to the numerous and

diverse species that use its burrows. Franz (1986) listed

19 species of amphibians and reptiles that used the SL bur-

rows. The most recent state-wide list of species associated

with Gopherus burrows consists of 60 vertebrates (Amphibia,

Reptilia, Aves, and Mammalia) and 302 invertebrate species









(Gastropoda, Malacostraca, Arachnida, Chilopoda, Diplopoda,

and Insecta) (Jackson and Milstrey, 1989).

Besides using burrows as shelter, I suspect Podomys

takes advantage of the invertebrate commensals and visitors.

My captive mice readily accepted live crickets and meal-

worms. Milstrey (1987) described Podomys eating blood and

body fluids of engorged ticks (Ornithodoros turicata) that

parasitize Gopherus and Rana areolata.

Florida mice are presumably preyed upon by birds,

foxes, bobcats, and raccoons (Layne, 1978; Maehr and Brady,

1986; Wassmer et al., 1988). I suspect that in the high

pine snakes are major predators, although predation by

snakes has not been documented. I occasionally saw coral

snakes (M. fulvius) near burrows, but omitted the species

from consideration because coral snakes do not prey upon

mammals (Jackson and Franz, 1981). Mushinsky (1987) sum-

marized predation records for snakes; some of the species

known to take mammals include Coluber constrictor, Drymar-

chon corais, Elaphe obsoleta, and Pituophis melanoleucus.

Several species of snakes utilize tortoise burrows at Ordway

and must be considered potential predators of Podomys,

including C. constrictor, E. quttata, P. melanoleucus,

Masticophis flagellum, D. corais, Sistrurus miliarius, and

Crotalus adamanteus (Franz, 1986; Timmerman, 1989). In

particular, eastern diamondbacks are known to take Peromys-

cus, and Timmerman (1989) considered Podomys a potential









prey item. During my time at Ordway, I saw only one dia-

mondback in the high pine (an adult female near BP burrow

#319, 6 November 1988). Timmerman (1989) examined habitats

used by eight eastern diamondbacks at Ordway from 1985 to

1989. His snakes used high pine and old field habitats less

than expected (approximately 13% of all observations) al-

though these two habitats comprised more than 60% of the

area available. The diamondbacks spent little time under-

ground, and only 16% of the burrow locations were in Gopher-

us burrows. However, as Timmerman pointed out, 75% of the

locations were in dens of the armadillo, which frequently

takes over old Gopherus burrows. Tortoise burrows are used

as winter refugia and, in the fall, as parturition sites,

leading to the suggestion that Podomys might be important in

the diet of young diamondbacks (Timmerman, 1989).

Mortality rates of Podomys due to disease and parasites

are unknown. Thirty-two species of endo- and exoparasites

of Podomys have been reported (Layne, 1963, 1968a; Lichten-

fels, 1970; Whitaker, 1968). Milstrey (1984, 1987) sum-

marized recent studies of the soft tick Ornithodorus, a

known carrier of relapsing fever that feeds on residents of

gopher burrows, but, as he points out, the relationships

between the vertebrates and invertebrates of these burrow

systems are still poorly understood.









Life History

I examined demographic data for my largest sample, the

255 P. floridanus marked at the SL burrows, August 1983 to

November 1988. Of this total, 52 females and 44 males were

marked in 1983. Eighty-one (38 females, 43 males)of the 255

(32%) were captured only once. I calculated minimum trap-

pability ((number of captures 2)/(possible captures -2))

for animals captured three or more times (Hilborn et al.,

1976).

Individual trappability ranged from 14 to 100%. Aver-

age trappability was 58% for 27 mice in 1983, 55% for n = 17

in 1984, 58% for n = 16 in 1985, 62% for n = 16 in 1986, 48%

for n = 13 in 1987, and 60% for n = 24 in 1988. The average

for all five years was 57%. According to Hilborn et al.

(1976), when trappability = < 50%, estimates of minimum num-

bers of individuals (MNI) by direct enumeration will under-

estimate actual population size; estimates become more

reliable as trappability increases. Trappability on grids

was lower than on burrows. For example, individual trap-

pability on the SL grid ranges from 38 to 50%.

The reproduction, growth, and development of Podomys

were described by Layne (1966). He demonstrated that sea-

sonal distribution of pregnancies in a wild population in

Alachua County was bimodal, with the major peak occurring

between August and November. At SL, I classified young

animals as juveniles if the gray and white juvenile pelage







35

was still present, and as subadults if the dorsum was gray

with gold and agouti hairs appearing along sides and flanks

(minimum weights of subadults were approximately 25 g).

Numbers of young marked (Fig. 5) and numbers of reproductive

adults (Fig. 6) at SL indicate that seasonality of reproduc-

tion at Ordway also peaked in the late summer and fall,

although juveniles and reproductive adults might appear in

small numbers at other times of the year (Layne, 1966).

Juveniles usually were first captured at the mouths of

burrows at approximately three weeks of age. At this point

the young were still completely gray and white and weighed

approximately 9 to 14 g (Layne, 1966). In some cases young

appeared at burrows with females (and sometimes males) who

were long-term residents, but I had no way of determining

parentage of some juveniles and almost all subadults. This,

and the fact that many juveniles disappeared before comple-

tion of adult molt, suggests that both pre- and post-

reproductive dispersal occur.

Captive animals breed year-round (Dice, 1954; Layne,

1966; Drickamer and Vestal, 1973). In captivity mean litter

size (1.7-2.6) is slightly smaller than that reported (3.1)

for wild-conceived litters (Dice, 1954; Layne, 1966; Rood,

1966; Drickamer and Vestal, 1973; Glazier, 1985). Maternal

care in captivity consisted of creating nests, plugging

entrances to nest boxes, nursing and grooming young, and

herding juveniles into the nest, as described previously by










an
I-
z
0






I-


z
2-


M A M J J A S O N


MONTH


-0-"


M A M J J A S O N


MONTH


Fig. 5. Numbers of juveniles (dots) and subadults
(circles) marked on the SL sandhill, expressed as
minimum numbers of individuals/number of
trapnights (top) and as numbers of individuals
(bottom).


1.2

1.0

0.8

0.6

0.4

0.2

0.0




























F M A M J JA S O N
MONTH


F M A M J J
MONTH


---t-0--


1984
1985


-**...." 1986


---+-0-













---


1987
1988















1984
1985
1986
1987
1988


Fig. 6. Reproductive data for Podomvs trapped on the SL
sandhill, 1983-1988: the proportion of adult
females who were pregnant (top) and the proportion
of males who were scrotal (bottom) are shown.









Layne (1966, 1968b). In my captive colony 8 females and 5

males produced 54 young in 24 litters; 11 litters were born

to a single pair. Litter size ranged from 1 to 4 (mean =

2.25). The distribution of births was: 8 in September; 4 in

July; 3 in October; 2 in March, August, and December; 1 in

January, February, and November. Four neonates have not

been sexed at this time; the remaining young consisted of 24

females, 19 males, and 7 infants who died before gender was

recognized.

Longevity of Podomys in the wild is unknown. Keim and

Stout (1987) reported that a male captured at approximately

three weeks of age lived in captivity for 7 years and 4

months. At SL, 12 females and 10 males (8.6% of the animals

marked) were present for more than 360 days (Table 3).

Seven females and one male were first marked as juveniles;

20 individuals were first found on the south side of SL and

two were marked on the north and later migrated south. In

captivity, two- and three-year-olds are continuing to repro-

duce.









TABLE 3. Longevity records exceeding 360 days for Podomvs
floridanus at SL. ID is the identification number, YR =
year marked, A = age when marked (Juvenile, Subadult, or
Adult), and P = persistence (in days) as indicated by both
trapping methods. Asterisk indicates trap mortality.


Females

ID YR

43 83

55 83

65 83

23 84

30 84

61 86

10 87

19 87

26 87

47 87

58 87

64 87


P

445

409

649

467

403

360

588

385

554

547

392

504


Males

ID YR

39 83

48 83

63 83

11 84

32 84

98 85

80 86

98B 86

42 87

54 87


P

905

570

410

920

620

401*

515

500

457

412













HOME RANGE


The concept of home range most widely accepted by

mammalogists is that defined by Burt (1943:351) as "that

area traversed by the individual in its normal activities of

food gathering, mating, and caring for young. Occasional

sallies outside the area should not be considered as

in part of the home range. The home range need not cover

the same area during the life of the individual." The

concept of territoriality, on the other hand, involves

domination by either overt or covert social interaction. As

defined by Kaufmann (1983:9), a territory is "a fixed por-

tion of an individual's or group's range in which it has

priority of access to one or more critical resources over

others which have priority elsewhere or at another time."

Among peromyscines, there is considerable variation in

descriptions of both home range sizes and degrees of ter-

ritoriality (Eisenberg, 1968; Stickel, 1968). For P. flori-

danus, home range and territoriality have not been des-

cribed. I examined the size of home ranges on the Ordway

sandhills with three methods: fluorescent powder, trapping

on grids, and trapping at burrows.










Fluorescent Powder

The use of fluorescent powder to track small rodents

and to determine movement patterns was introduced in the

sandhills of Nebraska (Lemen and Freeman, 1985, 1986). I

applied this technique to examine size of nightly activity

areas of Podomys on the SL sandhill.

I powdered five animals on 25 May 1985, three on 14

July 1985, and three on 15 June 1987. The first eight ani-

mals were trapped at burrows and held until 2100 EST the

following evening; the last three were removed from traps on

the grid at 0230 EST, powdered, and released immediately.

Each animal was placed into a plastic ziplock bag with a

fine, nontoxic fluorescent powder (green, pink, or yellow),

gently shaken until completely covered, and released at the

point of capture. At a later time, I returned with the

fluorescent light and marked the powder trails with forestry

flags. Trails then were mapped using a tape measure and

compass.

The mice showed no ill effects from this technique by

loss in weight or other signs of poor condition. One female

was so little traumatized by the procedure that she caught

and consumed a moth immediately after her release. However,

with the exception of the moth incident, I did not obtain

additional information regarding feeding.

Powdering was of some use in that it showed animals

utilizing fallen trees and branches as runways, as I had







42

observed in escape responses during trapping. Douglass and

Reinert (1982) showed that utilization of fallen logs by

Peromyscus leucopus was not an artifact of trapping but part

of the animals' normal movements. Additional evidence of

climbing was indicated by the powder trail of an adult male

(#39), which extended 36 cm up a turkey oak snag.

The powder indicated only one case of social interac-

tion. On 15 June 1987 females 10 and 47 were caught in the

same trap. Female 47 was powdered, released, and recaptured

later that morning. Again, she was found in the same trap

with #10, who also had become covered with powder. Female

47 was first marked as an adult on the SL grid on 19 April;

she remained at the grid and surrounding burrows until at

least October 1988. Number 10 was a younger female marked

as a juvenile on 12 April at a burrow located on the grid;

by September she had moved to an area west of the grid.

Possibly #10 was a daughter of #47. Use of radioisotopes,

DNA fingerprinting, and other techniques would be more

appropriate and efficient methods to examine family rela-

tionships and dispersal of young Podomys.

I Distances traveled by mice, as indicated by powder

trails, are shown in Table 4. Trails for two animals were

very short (< 10 m), but I believed that a single night's

excursion exceeded 10 m. By trapping on 14 July 1985 and 15

June 1987, I discovered powdered mice at locations to which

no powder trails led. Thus, the powder trail showed only a










TABLE 4. Activity data from ten Podomys dusted with fluo-
rescent powder at SL. ID indicates sex and identification
number of mouse; R indicates the burrow or grid station at
which an animal was caught and released; ORI indicates
initial direction from release point (in degrees); DIST is
the greatest distance (in meters) from R as indicated by
fluorescent powder; DEST is the burrow or grid station at
which an animal was found the following morning. The actual


distance


from R to DEST is shown in parentheses.


DATE

25 May 85

25 May 85

14 Jul 85

25 May 85

15 Jun 87

25 May 85

14 Jul 85

14 Jul 85

15 Jun 85

15 Jun 85


R

35

13

81

58

6,2

10

81

22

9,4

9,1


ORI

222

125

302


DIST

5.5

11.9


9.9


70 17.7

47 20.0


162

140

108

273

250


15.2

28.0

19.2

2.0

15.0


DEST


71 (36.0)


6,0

10


15 (45.0)

23 (15.0)

9,5 (10.0)


49 281 11.5


ID

F23

F30


F39

F47

Mil

M32

M39

M49

M54


M98 25 May 85









part of the night's activities. Because the fluorescent

powder yielded so little information regarding home range,

feeding behavior, or social activity, I concluded that it

was of little use with Podomys at Ordway.



Trapping on Grids

Results of trapping on the three grids are shown on

Fig. 7, which includes data from all animals, including

transients caught only once or twice. A total of 18 animals

was trapped on the AC grids. I calculated persistence on

the grid (P,) as the number of days from first to last

capture. Time spent by each individual on the grid is

represented by the horizontal bars; counting bars vertically

indicates the size of the known population per month. The

largest population occurred on the SL grid in the spring of

1987 and 1988, when there were 8 individuals or 8 ani-

mals/ha. Excluding animals caught three times or less, mean

persistence time on this grid was 195.9 days for females and

137.2 days for males.

For 20 mice captured four or more times, I plotted the

capture locations and number of captures. To calculate the

home range area, I used the Exclusive Boundary Strip Method

(Stickel, 1954), in which I mapped the capture sites and

drew a boundary strip (equal to half the distance between

traps). I selected this approach because Stickel (1954)

compared the exclusive boundary strip, inclusive boundary









M J J A S O M A M J J S N

ACI

TN 600 200 300 300 300 300 300 300 300 300 300 300


(21)
(75)


(3)
(21)
(21)
(21)
(21)


M94
M95
M96*


F85*


ACII


S(1)


300 200 300 300


300 300 300 300 300


F88
M89*
M92


(33)


M86


Fig. 7. Trapping data for Podomys on the ACI, ACII, and SL
grids. ID indicates sex and identification
number; asterisks indicate animals first captured
as juveniles; TN indicates trapnights per month;
and persistence on grid in days is shown in
parentheses.


1987


1988







1988


A M J J A S O M A M J J A S N

SL

Tr300 600 100 300 300 200 300 300 300 300 300 300 300


F10
F18*_
F26*
F47*
F64


(36)
(449)
(517)
(364)
(24)
(91)
(148)


M11


M16
M19
M49
M54
M42
M98
M58


(381)
(289)
(24)
(1)
(1)
(74)
(2)
(1)
(76)
(44)
(74)


M102


(61)
(1)


Fig. 7-- continued


1987










strip, and minimum area methods of calculating home range

area and she determined that the minimum area method under-

estimated home range size by 64%, and the exclusive and

inclusive boundary methods overestimated home range size by

2%, and 17%, respectively.

On ACII only one individual (female 87 on Fig. 7) was

captured at least four times. However, her last two cap-

tures occurred across the grid from the first three, and

this pattern suggests dispersal rather than a large home

range, so I did not include her in the analysis. Home range

data are summarized on Tables 5 and 6. Three animals were

marked on the grids as juveniles (gray pelage) and died or

dispersed off the grid at approximately 60 days of age. Two

more animals (female 26 and male 54 at SL) marked as juven-

iles did not leave the grid. The male shifted his home

range, but I could not distinguish between juvenile and

adult home ranges for female 26, who subsequently appeared

to have an activity area larger than that of other females.

Seventeen adults trapped on the two grids (ACI and SL) had

an average home range of 804 m2. In general, males (mean =

910 m2) appeared to have larger home ranges than females

(mean = 683 m2), but the difference was not significant

(Mann-Whitney test at p = 0.01).

To determine whether ranges of females and males over-

lapped, I plotted home ranges of Podomys caught during May

1987 on ACI and May 1987, September 1987, and May 1988 on SL









TABLE 5. Home range data for 20 Podomys trapped on two
grids. Id is the identification number, N is the number of
captures, PG is the persistence on the grid in days, A is
the area of the home range in m2 for juveniles (A,) and
adults (AA).


Grid Id Sex NG PG A A


ACI 89 F 6 21 700

ACI 91 F 7 75 500

ACI 96 F 4 21 400

ACI 94 F 6 21 916

ACI 95 M 5 21 750

ACI 96 M 5 21 800

SL 61 F 5 24 1000

SL 10 F 6 42 500

SL 18 F 6 36 600

SL 26 F 17 449 1350

SL 47 F 12 517 300

SL 54 F 6 74 300

SL 42 M 8 289 1850

SL 49 M 10 91 1200

SL 54 M 16 381 1093 200

SL 16 M 9 148 800

SL 53 M 6 76 600

SL 56 M 7 44 500

SL 59 M 8 74 1000

SL 102 M 10 156 400







49

TABLE 6. Capture data for 36 Podomys at Anderson-Cue. ID
indicates sex and identification number; YR is year of first
capture; AGE is age at first capture (juvenile, subadult, or
adult); N is number of captures at grids (NG), burrows (N,),
and total (NT); P is persistence, in days, on grids (PG),
burrows (P,), and total (PT). Asterisks indicate trap
mortality.


ID YR AGE NB PB NG PG NT PT


F83 88 J 1 1 1 1

F84 88 J 3 3 3 3*

F85 88 J 1 1 1 1

F86 88 A 7 109 7 109

F87 88 SA 2 2 5 33 7 33

F88 87 A 1 1 3 3 4 9

F89 87 A 6 21 6 21

F90 87 SA 1 1 1 1

F91 87 SA 7 75 7 75

F92 87 A 1 1 1 1

F93 87 A 1 1 1 1

F94 87 A 7 164 1 1 8 164

F95 87 A 1 1 3 3 4 21

F96 87 A 1 1 4 21 5 40

F97 87 A 9 171 3 3 12 193

F98 87 J 1 1 1 1

F99 87 A 1 1 1 1

F100 87 SA 4 124 4 124









TABLE 6---continued


ID YR AGE N P, NG PG NT PT


M80

M84

M85

M86

M87

M40

M88

M89

M90

M92

M93

M94

M95

M96

M97

M98

M99

M100


SA

A

SA

SA

SA

A

J

J

J

A

SA

A

A

J

A

?

SA

A


216

39

101



2

1

1



1

48


2 106


150

1

1

1


216

39

101

1

2

1

1

1

1

48

5*

106

21

21

150

1

21

1







51

(Fig. 8), the three months during which populations appeared

to be the largest (Fig. 7). Generally, ranges of males and

females overlapped, but ranges of adult females did not

(except occasionally with juveniles or young adults), a

pattern which suggests intra-sexual territoriality (Kau-

fmann, 1983). A female juvenile (female 18) overlapped with

the ranges of an adult of each sex (Fig. 8A). Female 26,

the one with the unusually large range, overlapped somewhat

with female 61 in September 1987. The only males that

overlapped were three males at SL (Fig. 8C), two of which

possibly were siblings.



Trapping at Burrows

With the data from trapping at burrows, I estimated

home ranges by mapping movements of adults trapped more than

four times in a year on AC and SL, using the exclusive

boundary method (Stickel, 1954) as I did for grids. For

burrows on the perimeter of the study site, I plotted a

boundary of 10 m (which I considered a conservative estimate

of distance moved beyond the burrow). All capture points

were weighted equally, but in a few instances I omitted

points that suggested unusual forays outside the usual home

range of the animal. I then measured the home range area

with a Lasico Compensating Planimeter.

Seventy-five home ranges for 62 animals who fit the

above criteria, including three females on AC, are shown in












A






* U

U I


* ,\ S I


I i.


* LJ_ I Ir
S
*~ U, *


I,.


9I I~ -


S,


0i I
*
* .

-J


B rr


I- I L

L.


o *


& 0 0 a 0



* t


/ /'


*



L--J1


1- I
0 U I I 11 S


Fig. 8. Home ranges of Podomys trapped on the ACI grid in
May 1987 (A) (n = 6 mice, 600 trapnights) and on
the SL grid in May 1987 (b) (n = 8, 600 trap-
nights), May 1988 (c) (n = 9, 300 trapnights), and
September 1987 (D) (n = 7, 300 trapnights). Dots
represent the outer boundary of grid. Solid lines
represent home ranges of females, dotted lines
those of males.


IJi
*.'
U. _









Fig. 9. Home ranges for adult males frequently overlapped,

but those of females did not; two SL females who appear to

overlap in 1988 actually were separated temporally. This

separation of home ranges, similar to that observed on

grids, suggests intra-sexual territoriality. Animals who

survived more than one year (e.g., SL male 11 in 1985-87)

showed fidelity to the same general area every year. Home

ranges varied in size from 210 m2 (for SL female 26, cap-

tured at a single burrow 6 times in 1987) to 11007 m2 (for

SL male 11 in 1986). The average home range size for 35

females was 2601 m2 (SD = 1315.1) and for 40 males was 4042

m2 (SD = 2269.4); home range size of the two sexes differed

significantly (Mann-Whitney U = 2.95, p = 0.01). For the SL

data I also compared areas in 1983-85 with those of 1986-88

to see whether home range size changed after the prescribed

burn in 1985, but I found no significant difference.

Capture data for the 62 mice whose home ranges were

mapped are shown on Table 7. Females 86, 94, and 97 (1987)

were from AC; the remaining animals were on SL. Each animal

was trapped from 5 to 22 times during the year in question,

and almost all mice utilized more than one burrow (mean

number of burrows = 3.8). Estimates of persistence (P,)

were calculated by counting the number of days between first

and last captures spent by the animal in the area shown on

the figure. These estimates of persistence are not com-

parable across years because trapping effort varied each































Fig. 9. Annual home ranges of 62 Podomys on the SL
sandhill, 1983-1988. Dots represent tortoise
burrows where mice were trapped. Solid lines
indicate home ranges of females, dashed line for
males; a) 1983, 52 burrows; b) 1984, 61 burrows;
c) 1985, 89 burrows; d) 1986, 103 burrows;
e) 1987, 131 burrows; f) 1988, 136 burrows. Bar
represents 20 m.



























--


0
S


*


53f-
.
* i '.
o S
* I


33


N


29
( *

S



48

S \



27


Fig. 9a.


49


I


j


4* -




























27,
SI


*4
*


*\ \
2
* '
I




23


* S
0




0



/
/
0


,- 48


25


38


29
*


3 9 .. "
39
85


Fig. 9b.


43


3






34

















100


*0
U ~


23


/ .


0
'0
I
I


97


39



0 0



-- 1 U


*.

* */
* --

/ 98
// 98


/ *


100


---39


Fig. 9c.


* 6

















97 .---
.* 995






11 --
S. 76






S73



11'
9 / -
.. 7 .,,




-*--
/. *.. \ ,...

A, '.' / *


0 98
*l / t ##

0 .





/



9 *
S80 0


Fig. 9d.


















17

65 --'




S 's *,
I I



S 68
47
/ .

*/4



S 7-* S. 0j


0 /
0 r




**26 *
----)~ri ~, \rr-

-----


* I*
* I *




72 --*


S *

* 9 8

.-'

** e l

57 *J
*
* 0


Fig. 9e.


* S


* *
**


. 80l -































S S
0
0


37


* 36
-
\ I \


49


-- z -I~


* *


20


10


Fig. 9f.


34


37


/
I
I
I

0

6 K


.
* /


102 '







61

TABLE 7. Capture data for the 62 adults whose home ranges
are shown on Fig. 9. ID is the sex and identification
number of each mouse; YEAR is the year for which the home
range was calculated; N is the number of captures, P, is
persistence (in days); # is the number of burrows where
each animal was caught; and asterisks denote animals present
when trapping ceased in November 1988.


ID YEAR N, P, #B


F5 1983 5 93 2

F8 6 71 3

F14 6 71 2

F27 5 114 2

F49 5 117 4

M29 5 79 2

M33 5 79 2

M47 5 99 2

M48 5 100 1

M53 5 79 2

F23 1984 8 125 2

F29 11 139 5

F34 5 56 2

F38 5 49 2

F43 15 222 3

M25 6 135 3

M27 5 61 3

M28 5 63 2


171 5


M39









TABLE 7--continued


ID YEAR N3 Ps #3


M48

M85

F23

F39

F30

F97

F100

M11

M32

M39

M98

M100

F89

F93

F94

F95

M11

M72

M73

M76

M80

M88

M97


1985


211

99

212

84

183

185

265

253

183

265

265

100

49

36

103

107

253

68

79

133

103

259

101


1986










TABLE 7--continued


ID YEAR N, P, #


M98

F26

F47

F61

F68

F72

F86

F94

F97

M11

M16

M17

M42

M57

M65

M80

M98

F10

F26

F36

F37

F46

F47


1987


171

78

113

243

170

125

109

164

171

143

233

193

31

242

213

201

218

227*

193

94

121

151

193


1988









TABLE 7--continued


ID YEAR N, PB #B


F49

F52

M20

M22

M25

M34

M37

M42

M51

M102


148*

151

151

151

132

104

148*

198*

151

198*









year; nor do they represent longevity because captures of

juveniles and subadults (or captures as adults in years

other than those shown) are excluded. The P, values are

useful as minimal estimates of persistence on the sandhill.


Conclusions

Fluorescent powder was not a useful method of determin-

ing the home range of Podomvs on Ordway. Powder trails did

not indicate the complete area traveled by mice even during

one night. Although some data regarding substrate use or

social interactions were gathered, considerable time and

effort were required. The technique proved successful with

smaller Peromyscus (Lemen and Freeman, 1985, 1986), so I

suspect the failure at Ordway was not related to the body

size or biology of Podomys, but possibly to density or

structure of vegetation.

Trapping on grids provided estimates of home range size

and persistence, as well as demonstrating the exclusivity of

home ranges of adult females. There also was evidence that

males had larger home ranges that occasionally overlapped

with each other and with females. However, estimates of

home range areas obtained from trapping at burrows suggest

that estimates from grids were too small. Evidently the

"standard" mammal-trapping grid (10000 m2 with 10-m trap

intervals) was insufficient to measure home ranges of Podo-

mvs in sandhills. Doubling the trap interval to 20 m would







66

produce a 10 row by 10 column grid covering 40000 m2, which

would encompass mean home ranges estimated at burrows for

females (2601 m2) and males (4042 m2), might be more useful.

No fewer than 100 trap stations should be used. However,

one should keep in mind that increasing grid size on some

sandhills will spread the grid into neighboring habitats,

and that trap success on the grid will be smaller than on

the burrows.

Trapping at burrows produced larger estimates of home

range size than I expected, but the means are not unreason-

ably large when compared with other values reported for

Peromyscus. Stickel (1968) reported that home ranges of

Peromyscus varied from 0.1 to 10 acres. Hoffmeister (1981)

reported mean home ranges for female (8290 m2 +/- 2688) and

male (10465 m2 +/- 4043) P. truei in pinon-juniper habitat,

also using the exclusive boundary strip method. Although

the range of home areas of Podomys varies considerably,

further research might show that my means are under-

estimates, based as they are on many individuals located on

the edges of my study sites. I also have no measure of

vertical habitat use; for example; it would be interesting

to determine whether the mice occupy a larger home range

volume in years when acorns are produced.

In analyses of both burrow and grid data, I omitted

animals that were recaptured less than three times. A pos-

sible consequence of this approach is that several animals







67

might occupy the same space. However, the fact that both

sets of data show abutment of home ranges of adult females

seems good evidence of mutually exclusive home ranges, which

suggests that females might be territorial.













DIET


Field Observations

Almost nothing is known about the natural diet of

Podomys. Merriam (1890) reported an observation that mice

in the southeast ate the seeds of "scrub-palmettoes." Layne

(1970) and Humphrey et al. (1985) suggested that acorns

might be a major food source during mating years, based on

the association of P. floridanus with habitats with Q.

laevis and other oaks. Milstrey (1987) reported that captive

Podomys ate engorged soft ticks (Ornithodoros turicata

americanus) that parasitize gopher frogs and tortoises,

raising the possibility that these mice prey on other resi-

dents of Gopherus burrows. Other foods presumably include

insects, seeds, nuts, fungi, and other plant material

(Layne, 1978).

Five field observations at SL provided additional

dietary information. An adult male caught and released on

the grid (5 May 1988) readily ate a cricket (Gryllinae) I

offered to him. On 15 June 1987 at approximately 0245 EST,

an adult female (who had been trapped, powdered, and re-

leased) caught a small moth and consumed all but the wings.

On 9 May, an adult female just released from a trap ate a









young shoot of Smilax auriculatus. On 20 July 1987, at

burrow #15, I also observed an adult male take a fruit of

the flag pawpaw (Asimina incarna) (Jones, 1989). After

being released from the trap, this mouse climbed 30 cm up a

plant on the apron of the burrow, and knocked a ripe pawpaw

to the ground. He then ran down, picked up the pawpaw, and

carried it 11.4 m to burrow #71. In July 1988 another mouse

took a pawpaw, again at SL. A female released from a trap

at a burrow hid in a hole at the base of a turkey oak ap-

proximately 5 m NW of the burrow. In a few minutes she left

the hole, picked up a small pawpaw fruit (A. incarna), and

carried it back to the hole and nibbled on it.

Predation and dispersal of Asimina fruits have not been

well studied. Willson and Schemske (1980) reported opossums

and raccoons sampling unripe fruits of A. triloba in Il-

linois. Norman and Clayton (1986) suggested that small

mammals probably help disperse Asimina in Florida and re-

ported an observation of Gopherus eating ripe fruits. Human

consumption of A. incarna was reported by Bartram (1791).

Whether Podomys disperse these fruits, and whether pawpaws

are a significant source of food, remains unknown. Willson

and Schemske (1980) measured a caloric content of 4.5 kcal/g

for fruits of A. triloba in Illinois; fruits ranged in size

from 2.1 to 48 g dry weight. Fruit set for A. obovata, A.

Dvgmaea, and A. triloba is less than 10% (Willson and

Schemske, 1980; Norman and Clayton 1986). To the best of my









knowledge, nutritional values and fruit set have not been

reported for A. incarna, the most abundant pawpaw on Ordway.

However, pawpaws certainly are the largest fruits produced

in the high pine; surely they represent a significant addi-

tion to the summer diet of Podomys.



Laboratory Observations

The maintenance diet of captive animals consisted of

rodent chow (Wayne Rodent Blox) and water provided ad libi-

tem. This diet was supplemented with lettuce, carrots,

apples, strawberries, sunflower seeds, mixed bird seed,

oatmeal, rice, mealworms, crickets, and acorns (Q.

michauxi).

In an attempt to determine what mice might eat under

natural conditions, I occasionally offered animals fruits

and seeds collected at Ordway. All of these offerings were

eaten: acorns (Q. chapmanii, Q. geminata, Q. hemisphaerica,

Q. laevis, Q. myrtifolia, and Q. nira), pine seeds (P.

elliottii and P. palustris), blueberries (Vaccinium mvr-

sinites), deerberries (V. stamineum), gallberries (Ilex

qlabra), blackberries (Rubus arqutus), gopher apples (Lica-

nia michauxii), pawpaw fruits (A. incarna and A. yvcmaea),

flowers of Stillingia sylvatica, and seed pods of Crotalaria

rotundifolia and Galactia elliotti. Seeds and stems of

unidentified grass species were shredded and incorporated









into the cotton nesting material, and probably eaten as

well.

Predation on vertebrates appears limited to stressful

conditions. Eisenberg (unpubl. data) trapped an adult male

on SL in 1983 who detached the posterior part of a juvenile

red rat snake (Elaphe quttata) that was caught half way in

the trap. The mouse ate about 3.5 cm of the snake to the

bone. An adult male caught on AC in October 1987 ate the

viscera of a juvenile also caught in the trap. I also lost

a litter of three young in captivity on the only occasion

that I had a male and two pregnant females in the same

aquarium.

In general, feeding behavior follows that described for

Peromyscus by Eisenberg (1968), in which the animal crouches

and manipulates the food with its forepaws. Larger items

such as pawpaws are held against the substrate. Seed pods

were opened by grasping the pod vertically, resting one end

on the substrate, chewing off one end of the pod, and open-

ing the pod longitudinally along a suture. Larger foods,

such as turkey oak acorns, are dragged with the incisors;

smaller items are carried in the mouth. Some individuals

consistently took favored foods (such as acorns and sunflow-

er seeds) to the corners of the aquarium and covered them

with kitty litter; I considered this caching behavior. Food

items and remains also commonly are found underneath nests.









This ready acceptance of a wide variety of foods im-

plies that P. floridanus is an opportunistic feeder, like

Peromvscus. As King (1968) pointed out, we lack systematic

studies of taste discrimination and food preferences. For

Podomys we also need basic studies (stomach and fecal ana-

lyses) to learn more about the natural diet.



Acorn Preference Tests

I performed preference tests to determine whether

acorns of turkey oaks (Q. laevis) are preferred over acorns

of other species that are less abundant or produce fewer

acorns in the high pine. Such a preference might be a

partial explanation for the distribution of Podomys in

sandhills at Ordway.

For feeding tests, an individual was placed in an

aquarium with clean kitty litter, nesting material, water,

and rat chow at least 24 hours before beginning the test.

To start a trial the chow was removed and three bowls (each

containing five acorns of a single species) were added at

about 1900 hours. Each acorn was marked and weighed and I

measured its height and width with dial calipers. No effort

was made to ascertain that all acorns in a bowl were identi-

cal in size. Acorns that had weevil holes were not used,

and in a single trial all acorns either had caps or lacked

them. Approximately 12 hours later I removed the bowls,

acorns, and acorn fragments. I recorded whether acorns were










removed from bowls and whether they were opened and eaten,

opened and evidently not eaten, gnawed, or apparently un-

touched. I controlled for location effects by shifting the

relative positions of the three types of acorns in each

trial.

Two unexpected difficulties arose. The first was the

unpredictability of the acorn supply, which made it impos-

sible for me to run tests with identical species in 1988 and

1989. Additionally, the majority of acorns on the ground

already contained weevil larvae or were otherwise spoiled.

Secondly, I discovered that some of the acorns I tested were

spoiled (by insects or by mold) even though they lacked

holes. Possibly this spoilage is the reason that some hulls

were opened but the meat not removed. For this reason I

simplified analysis of results by examining whether acorns

were unopened or opened, regardless of whether any of the

meat appeared to have been removed. These data were then

ranked (rank 1 to the smallest number opened) and the Fried-

man test (Conover, 1980) was used to test the hypothesis

that species of acorns were opened in equal numbers.

In 1988, I concentrated on determining which acorns

were eaten by Podomys. I presented acorns from six species

of oaks (Chapman's (Q. chapmanii), live (Q. geminata),

turkey (Q. laevis), laurel (2. hemisphaerica), myrtle (Q.

mvrtifolia), and water oaks (Q. niara)) to four captive

animals. Five of these species belong to the red-oak group,









which generally contains three or four times more tannin

(Briggs and Smith, 1989) than species of the white-oak group

(which includes Q. chapmanii). Each mouse was presented

with turkey oak acorns in combination with acorns from two

other species; two to four trials were run per animal for a

total of 12 trials. Although the sample is inadequate for

statistical analysis, I noted that the mice opened acorns of

all species, and in all but one trial turkey oak acorns were

opened in the smallest numbers. I then planned to test

animals with a combination of acorns from Chapman's, live,

and turkey oaks, the most prevalent Quercus species on the

SL sandhill.

In 1989 Chapman's and live oak acorns were not avail-

able, so I gathered acorns from Q. laevis and two different

trees of Q. hemisphaerica, one from the old pasture near

Ross Lake Bridge and the second from a hammock past Ander-

son-Cue. I expected that Q. laevis, the predominant acorn-

producing species on SL, would be preferred. I tested nine

animals, two trials each. For analysis I used results of

the first trial only; there was no difference in ranks of

first and second trials for any of these mice. Results

indicated that acorns were not opened in equal numbers

(Friedman test, T = 26.75, significant at p = 0.01). When

the Friedman test indicates rejection of the null hypothe-

sis, multiple comparisons may be made to determine which

treatment is significantly different from the others







75

(Conover, 1980). For multiple comparisons at a significance

level of p = 0.01, acorns from Ross Lake (Q. hemisphaerica)

were opened significantly more often; differences between

the acorns from the hammock and from Q. laevis were not

significant. These results indicate not only that the mice

preferred laurel oak acorns over turkey oak acorns, but that

mice distinguished acorns from two different individuals of

Q. hemisphaerica.

The apparent preference for Q. hemisphaerica was sur-

prising in that its acorns contain more tannic acid than Q.

laevis (Harris and Skoog, 1980). The laurel acorns were

smaller than the turkey oaks', so possibly the preference is

due to differences in handling time. Another possibility is

selection according to chemical composition other than

tannin content. Halls (1977) reported that Q. laurifolia

(= Q. hemisphaerica) had higher fat and carbohydrate content

than Q. chapmanii and Q. incana; unfortunately, values for

Q. laevis were not listed. Possibly my results were some

artifact of using captive animals. However, in the only

published study of acorn preference in Peromyscus, Briggs

and Smith (1989) found that five P. leucopus captured in

habitats lacking oaks consumed equal amounts of acorns from

species of the red and white oak groups. Peromvscus caught

in areas containing oak trees selected acorns of Ouercus

species found in their habitat, independent of fat, protein,









and tannin content. These results lead me to believe that

there is some real difference between the acorns I used.

Except for smaller acorns that occasionally were split

in half, acorns were opened consistently at the hilum (basal

scar). Although acorns typically were carried by the point

at the distal end, the mice never chewed open the hull

there. For small, round acorns a neat incision was made

around the scar; on more elongate nuts (Q. geminata and 2.

laevis) the hull might be nibbled farther down the sides.

Caps, if present, were removed; there was no difference in

the numbers of capped and capless Q. laevis acorns opened.

Occasionally I found acorns opened in this manner on aprons

or near burrow entrances. Hulls of Q. laevis were found in

excavated burrows (Jones and Franz, in prep.) and E. Mil-

strey vacuumed remains of what appear to be Q. qeminata and

Q. laevis out of burrows on Roberts Ranch, Putnam County;

all of these hulls were opened in a manner consistent with

what I observed with captive animals. Small piles of emptied

hulls occasionally are found at the base of turkey oaks on

Ordway; however, it is possible that Glaucomys or birds

might make these piles. Of the 450 acorns presented in 30

trials to 13 mice, 66% were removed from bowls, whether

opened or not.

I avoided use of acorns that appeared spoiled because

mice did not appear to eat blackened nutmeats. I did not

test preferences of sound acorns vs. acorns with weevil









larvae. On one occasion an adult female immediately ate a

larva from an acorn opened but not eaten by another in-

dividual. Semel and Andersen (1988) suggested that such

differences in behavior might be due to mice being unable to

detect larvae in unopened acorns, or that individuals can

detect infestation but some individuals avoid infested nuts.

They thought that tooth marks on the hull and movement of

acorns might represent assessment.

Based on these results, I suggest that acorns of Q.

laevis are eaten but other species are preferred if avail-

able. Turkey oaks are an unreliable food supply. Umber

(1975) noted acorn production of Q. laevis was low and

variable on his study site in Citrus/Hernando counties, and

Kantola and Humphrey (in prep.) found that trees on low

slopes at Ordway produced significantly more acorns than

trees on higher slopes. Layne (in press) correlated the

relatively greater abundance of Podomys in scrub and scrubby

flatwoods with higher and more consistent acorn production

than in the high pine. Proximity to Q. geminata and other

acorn-producers might be a partial explanation for the

relatively higher and more constant population of Podomys at

SL than at AC and BP. If oak species in hammocks are more

reliable producers, the restriction of P. gossypinus and

Ochrotomys nuttali to hammocks at Ordway might be partly due

to the food supply. This leads to the hypothesis that

competition, possibly linked with other factors such as








78
predation, forces Podomys into the high pine when the other

rodents are present.



Supplemental Feeding Experiment

Are Podomys limited by food supplies? If food is a

limiting factor, mice should respond to extra food with an

increase in numbers, increased individual mass, and an

extended breeding season. A food limitation could partly

explain population sizes and distributions at Ordway. I

attempted an experiment of food supplementation using foods

favored by captive Podomys. The only supplementation ex-

periment that has been done with the Florida mouse was that

by Young (described in Young, 1983 and Young and Stout,

1986) on two grids in sand pine scrub in Orange County.

Other rodents responded to the additional food, but Podomys

rarely appeared on Young's grids and failed to establish a

permanent population during the experiment, although the

species had been abundant on both grids before the study

(Young, 1983). She concluded that Florida mouse populations

were limited by factors other than food.

My experimental design consisted of three grids, two on

the AC and one on the SL sandhill. Trapping procedure was

described under Home Range. The last grid (SLII) was set on

the SL sandhill west of Breezeway Pond. Prior trapping at

burrows in each of these areas indicated that mice were

present. In April, May, and June of 1988 I trapped SLI for








79

800 trapnights to determine how many nights of trapping were

needed to obtain all animals on the site. Consequently,

SLI, ACI, and ACII were trapped for three consecutive nights

per month for a total of 300 trapnights monthly. On ACI a

mixture of sunflower seeds, mixed bird seed, and oatmeal was

provided for one year in seven chick feeders fitted with

glass jars. To eliminate non-target species, plastic buck-

ets with two one-inch holes cut near the rims were upended

over the feeders and weighted down with a cinder block. I

checked feeders periodically and refilled as needed.

The results of this experiment are shown in Fig. 10.

Capture rates declined sharply on AC and were highly vari-

able on SL. I had intended to provide food on a second grid

(SLII, west of Breezeway Pond). However, 208 trapnights on

this grid and surrounding burrows in May, July, and August

resulted in the capture of one Podomys. The lack of success

on ACI led me to abandon SLII although I continued to pro-

vide food on ACI until the end of the 1988 trapping year.

On the AC sandhill I continued to trap mice in small numbers

at burrows west and east of ACI. Clearly, mice were not

obtaining the food or some other factor kept them out of the

grid area in spite of obtaining food.

Maximum trapping success for a single night was 9%, but

success was usually much lower. Additionally, I captured

non-target species, including Sceloporus undulatus, Coluber

constrictor, and Glaucomys volans. In spite of the low trap




















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82

success I attempted to estimate the persistence (i.e., the

time between first and last captures) of mice on the three

grids (Table 8). Because these were not closed grids,

disappearance of an animal might be due to death, dispersal,

or to not entering traps.







83

TABLE 8. Persistence data for animals on grids. Data
presented are minimum number known alive (MNA), mean +/- SD
days spent on a grid (maximum values in parentheses), and
total numbers of trapnights (TN).


Grid MNA PERSIST TN


ACI 12 16.00 +/- 20.86 (75) 4200

ACII 6 7.17 +/- 12.68 (33) 3000

SLI 24 122.75 +/- 155.7 (517) 4100













ANALYSES OF VEGETATION


Sampling Technique

Can the local distribution of Podomys be predicted on

the basis of vegetation structure or species composition?

Vegetation was sampled in circular quadrats (radius = 10 m,

area = 314.159 m2) set with tortoise burrows as center

points. In the absence of large numbers of trees, circular

quadrats are laid out easily using a center pole at the

burrow and a rotating radius line; forestry flags were used

to delineate the quadrat. Quadrat size was based on prelim-

inary estimates of activity areas obtained by dusting Podo-

mys with fluorescent powder. Although one animal ranged at

least 28 m from his burrow in one night (see Home Range

above), I decided a radius of 10 m was sufficient to contain

significant numbers of plants and small enough to allow me

to feasibly obtain counts and identifications of plant

species. When I began these samples in 1985 I was uncertain

that burrows were distributed randomly, so point-quarter

sampling seemed inappropriate. Without knowing the shape of

Podomys activity areas, I felt that line-intercept sampling

and belt transects would be less representative of areas

encountered by mice than circular quadrats.








85
Sandhills on the Ordway Preserve have been subjected to

logging, grazing, suppression of fire, and prescribed burn-

ing on different rotations. Cessation of fire, or the

frequency of prescribed burns and other activities, affect

plant species composition and density on sandhills (e.g.,

Myers, 1985; Gates and Tanner, 1988). To determine how

burrow use changes with successional changes in vegetation,

I examined quadrats at SL in 1985, 1987, and 1988. SL has

been burned and the pines bear turpentine scars. Trapping

effort was equal among these burrows within each year but

not among years. Initially 29 quadrats were examined; this

number was reduced to 27 after a large oak fell over one

burrow and a clearcut was performed at a burrow under a

powerline. To see how Podomys behaved at burrows on a

sandhill where logging occurred, I set quadrats at 15 bur-

rows at BP in 1988.

I selected burrows so as to include burrows of all

three classes of tortoise activity (active, inactive, or

old, as defined by Auffenberg and Franz, 1982). Within each

quadrat, I counted stems of all woody species (Table 9) in

three size categories (tree = dbh > 7.6 cm, sapling = height

> 1 m and dbh < 7.6 cm, and seedlings = height < 1 m). I

noted the presence or absence of woody debris, including

fallen branches or trees, or rotting logs. I estimated the

per cent ground cover of bare ground, wiregrass, and gopher

apple using five categories (I = 0-10%, II = 10-25%,









TABLE 9. Habitat variables measured in each quadrat. Prime
signs indicate variables used in discriminant analyses.


Variable (abbreviation)


Location (LOC)'
Year (YR)'

Burrow (BUR)'

Orientation (ORI)'


Sand pine (SP)'
Slash pine (HP)
Longleaf pine (LP)'

Loblolly pine (YP)
Bluejack oaks (BO)'
Turkey oaks (TO)'
Laurel oaks (LA)
Post oaks (PO)
Live oaks (LO)'
Persimmon (PER)'
Pawpaws (PAW)'
Rosemary (RO)'
Sumac (SUM)'

Blueberry (VAC)'
Debris (DEB)'


Description


BP or SL sandhill
1985, 1987, or 1988
Identification number

Compass reading away from
entrance

Total stem count (P. clausa)

(P. elliotti)
(P. palustris)

(P. taeda)
(Q. incana)
(Q. laevis)

(Q. laurifolia)
(Q. stellata)
(Q. geminata)
(D. virginiana)
(Asimina spp.)
(C. ericoides)
(R. copallina)

(V. myrsinites)
Presence or absence of fallen

logs or branches in each

quarter of quadrat (4 indi-
cates presence in all 4 quar-

ters)










TABLE 9--continued


Variable (abbreviation)


Bare ground (GRO)'






Bare ground (ZGRO)'




Wiregrass (WIR)'






Wiregrass (ZWIR)'




Gopher apple (GA)'






Gopher apple (EGA)'




Legumes (LEG)'


Herbs (HERB)'


Description


Average of categorical es-

timates of % of bare ground

coverage in each quarter of

quadrat

Summation of categorical

estimates of bare ground

coverage per quadrat
Average of categorical es-

timates of % of coverage by
wiregrass of each quarter of

quadrat

Summation of categorical

estimates of wiregrass cover-

age per quadrat
Average of categorical es-
timates of % coverage by
Licania of each quarter of
quadrat

Summation of categorical esti-

mates of Licania cover per

quadrat
Total number of species of

Fabaceae
Total number of all non-woody

species







88

III = 25-50%, IV = 50-75%, V = 75-100%). I grouped species

of grasses into five categories (I = Andropoqon spp.;

II = Panicum hemitonum, III = Panicum (Dicanthelium) spp.,

IV = Paspalum setaceum, V = Gymnopogon spp.) and sedges into

two ("small", such as Bulbostylis warei, and "large", such

as Cyperus retrorsus), but did not identify each species. I

identified and counted the number of herbaceous species

(excluding grasses and sedges), but I did not count stems.

All plant species (Appendix) were identified using the plant

list compiled for the preserve by David W. Hall and Richard

Franz.

I plotted a species-area curve (cumulative number of

species vs. cumulative number of samples) for the 1985 data

from SL; the curve levelled off after the fifteenth sample,

and I concluded that the number and size of quadrats were

adequate. I believe smaller sample areas would be inap-

propriate for two reasons: Hansen (1964) noted that only 8

of 24 burrows he examined had turkey oaks and pines within a

radius of approximately 305 cm around the burrow entrance,

so larger areas are required to represent trees in the

vicinity. Secondly, some quadrats I examined were located

at burrows with large aprons, which are disturbed areas that

might be favored by certain animals such as Tantilla relic-

ta, Eumeces eqregius, and Neoseps reynoldsi (Campbell and

Christman, 1982); it is unknown whether these species affect

plant distribution on the apron. I suspect that burrow




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81,9(56,7< 2) )/25,'$



MICROHABITAT USE BY PODOMYS FLORIDANUS IN THE HIGH
PINE LANDS OF PUTNAM COUNTY, FLORIDA
By
CHERI A. JONES
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
1990

Copyright 1990
by
Cheri Ann Jones

ACKNOWLEDGMENTS
I thank my chairman, J. F. Eisenberg, for providing
encouragement, advice, and equipment during this project.
Members of my committee (R. Franz, J. H. Kaufmann, R. A.
Kiltie, F. E. Putz, and J. G. Robinson) made useful comments
and suggestions regarding this paper. John Eisenberg and
Dick Franz bear responsibility for introducing me to the
Ordway Preserve, where I formed a strong attachment to
Florida mice, gophers, and sandhills. I also benefited from
discussions with C. A. Gates, J. I. Glendinning, J. N.
Layne, and members of the Eco-lunch seminar group in the
Department of Zoology.
I am grateful to the Florida Game and Fresh Water Fish
Commission for issuing permits allowing me to trap Florida
mice and to establish a captive colony. I borrowed aquaria,
traps, and other equipment from J. F. Anderson, L. S. Fink,
W. H. Kern, R. F. Labisky, C. A. Langtimm, F. Nordlie, P.
Ryschkewitsch, and C. A. Woods. The Florida Foundation
provided funds for additional traps, flags, and other field
supplies. I thank T. Perry, the preserve caretaker, for
providing information about the history of Ordway and for
pulling my truck out of the sand more frequently than I care
to remember. Help in caring for captive animals, counting
iii

plants, and other forms of assistance were provided by R. A.
Drury, J. Dudley, J. F. Eisenberg, K. Eisenberg, J. Ellis,
N. Etheridge, R. Franz, S. E. Franz, B. Furlow, A. Gannon,
M. K. Lacy, C. A. Langtimm, A. Mahler, M. Marquez, R. Mor¬
rison, C. Partin, L. Partin, J. R. Thomson, and W. W. Tim¬
merman. I thank R. Franz and D. W. Hall, who patiently
identified plant specimens and taught me much about the high
pine flora. J. Boucher, J. I. Glendinning and K. M. Portier
helped solve statistical and computer problems.
Burns on the Ordway-Swisher Preserve were conducted by
J. F. Eisenberg, R. Franz, T. Perry, and M. E. Sunquist.
These burns would be impossible without additional help; I
thank all those volunteers (including R. Burke, P. Carnell,
B. Charest, C. K. Dodd, Jr., R. A. Drury, S. E. Franz, C. A.
Gates, H. Gold, L. Hay-Smith, C. A. Langtimm, M. E. Ludlow,
R. Morrison, D. Pearson, F. E. Putz, G. W. Tanner, K. K.
Timmerman, D. D. Wright, and students from the departments
of Botany, Wildlife, and Zoology) who cheerfully incinerated
my study sites and other sandhills.
Finally, special thanks are due to my parents, Clyde
and Charlene Jones, and my grandparents, Ellis and Maye
Owen, who have provided constant encouragement and moral
support throughout my academic career. I also am grateful
to R. A. Drury, P. Foster-Turley, R. and S. E. Franz, and C.
A. Langtimm, who offered friendship and support during my
years in Gainesville.
IV

TABLE OF CONTENTS
ACKNOWLEDGMENTS iii
LIST OF TABLES vii
LIST OF FIGURES ix
ABSTRACT xi
INTRODUCTION 1
TAXONOMY AND SYSTEMATICS 4
DISTRIBUTION 7
MATERIALS AND METHODS 10
USE OF TORTOISE BURROWS 18
Dependence on Burrows 18
Description of Burrows 26
Podomvs and the Burrow Community 31
Life History 34
HOME RANGE 40
Fluorescent Powder 41
Trapping on Grids 44
Trapping at Burrows 51
Conclusions 65
DIET 68
Field Observations 68
Laboratory Observations 70
Acorn Preference Tests 72
Supplemental Feeding Experiment 78
ANALYSES OF VEGETATION 84
Sampling Technique 84
Results and Discussion 89
v

PRESCRIBED BURNS
100
Materials and Methods 100
Description of Procedure for Prescribed Burns . . 102
Results and Discussion 103
Conclusions 129
SUMMARY AND DISCUSSION 132
APPENDIX LIST OF PLANT SPECIES IDENTIFIED IN
VEGETATION SAMPLES 140
LITERATURE CITED 146
BIOGRAPHICAL SKETCH 159
vi

LIST OF TABLES
Table Page
1 Numbers of captures of non-target species in
traps at burrows (22,085 trapnights) and on
grids (11,300 trapnights) 24
2 Data describing distribution of burrows on SL
(top) and AC (bottom) with the probabilities
expected from the Poisson distribution (with
means of 1.5 and 1.7 burrows per square) 30
3 Longevity records exceeding 360 days for
Podomys floridanus at SL 39
4 Activity data from ten Podomys dusted with
fluorescent powder at SL 43
5 Home range data for 20 Podomys trapped on two
grids 48
6 Capture data for 36 Podomys at Anderson-Cue ... 49
7 Capture data for the 62 adults whose home
ranges are shown on Fig. 9 61
8 Persistence data for animals on grids 83
9 Habitat variables measured in each quadrat .... 86
10 Frequency (% = percent of quadrats in which
a species occurred) and density (# = average
number of stems per 314 m2 quadrat) of woody
species identified at SL 90
11 Frequency (% = percent of quadrats in which
a species occurred) and density (# = average
number of stems per 314 m2 quadrat) of woody
plant species identified at BP in 1988 92
12 Habitat profile of Podomys at all BP and SL
burrows where vegetation was sampled 94
vii

13
Number (%) of vegetation samples reassigned
according to discriminant function analysis
to two categories of total number of captures
(ALL) o f Podomvs 96
14 Burrows on burned sandhills Ill
15 Burrows on unburned sandhills 112
16 Comparison of capture indices for Podomvs
at burrows on burned and unburned sandhills ... 113
17 Comparison of capture indices for Podomvs at
burrows on 3 burned sandhills 115
18 Survival of Podomvs following three prescribed
burns on the Ordway Preserve 120
19 Comparison of trapping effort on unburned and
burned sandhills, where TN is the total number
of trapnights and trapping effort is expressed
as trapnight/capture/ha (TN/CAP/HA) 122
20 Residence of young Podomvs on the SL sandhill,
where N is the total number of young marked
and R is the percentage of marked young that
became residents on that sandhill 123
21 Responses of Peromvscus species to fire 125
Vlll

LIST OF FIGURES
Figure Page
1 Distribution of Podomvs floridanus 8
2 Distribution of Podomvs on the Ordway/Swisher
Preserve 14
3 Trapping success (%) and minimum numbers of
individuals (MNI) (adults and subadults) of
Podomvs at burrows on the Smith Lake Sandhill • . 22
4 Trapping success (expressed as the total num¬
ber of captures/adjusted trapnights) for Podo¬
mvs on the AC (top) and SL (bottom) grids .... 27
5 Numbers of juveniles (dots) and subadults
(circles) marked on the SL sandhill, expressed
as minimum numbers of individuals/number of
trapnights (top) and as numbers of individuals
(bottom) 36
6 Reproductive data for Podomvs trapped on the
SL sandhill, 1983-1988: the proportion of
adult females who were pregnant (top) and the
proportion of males who were scrotal (bottom)
are shown 37
7 Trapping data for Podomvs on the ACI, ACII,
and SL grids 45
8 Home ranges of Podomvs trapped on the ACI grid
in May 1987 52
9 Annual home ranges of 62 Podomvs on the SL sand¬
hill, 1983-1988 55
10 Minimum numbers of individuals (MNI) on the ACI,
ACII, and SL grids 81
11 Apparent densities (expressed as minimum numbers
of individuals/ha) of Podomvs on the burned and
unburned areas of the Blue Pond sandhill 105

12 Apparent densities (expressed as minimum numbers
of individuals/ha) of Podomys on the burned and
unburned areas of the Longleaf Pine sandhill . .
13 Apparent densities (expressed as minimum numbers
of individual/ha) of Podomys on the burned and
unburned areas of the Smith Lake Sandhill . . .
14 Home ranges of eight Podomys on the SL sandhill
before (stippled) and after (hatched) the 1985
fire
107
109
117
x

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
MICROHABITAT USE BY PODOMYS FLORIDANUS IN THE HIGH
PINE LANDS OF PUTNAM COUNTY, FLORIDA
By
Cheri A. Jones
May 1990
Chairman: John F. Eisenberg
Major Department: Zoology
The purpose of this project was to examine the use of
tortoise burrows (Gopherus Polyphemus) by the Florida mouse
(Podomvs floridanus) in high pine (longleaf pine sandhills)
on the Katharine Ordway Preserve/Swisher Memorial Sanctuary
in Putnam County, Florida. Mark-and-recapture techniques
were used at tortoise burrows and on grids (10000 m2) in
order to describe burrow use by mice and to measure home
ranges. I also performed a food supplementation experiment
to test the hypothesis that population sizes were food-
limited. Finally, I examined how local distributions of
Florida mice were correlated with vegetation structure and
affected by prescribed burns.
Podomvs floridanus are commensals of Gopherus in high
pine habitats at Ordway, as indicated by trapping success,
escape responses, burrow excavations, and observations of
xi

mice in the field. In five years of trapping at burrows on
a single sandhill, 255 mice were marked. Of this total, 32%
were captured only once and 8.6% survived for 360-920 days.
Reproductive activity was concentrated in August, September,
and October. Florida mice appeared to be opportunistic
feeders, accepting a wide variety of fruits, seeds, and
insects. Populations appeared to be limited by factors
other than food. Mean annual home ranges were 2601 m2 for
adult females, whose ranges were mutually exclusive. Home
ranges of adult males were significantly larger (4042 m2)
and overlapped with those of other adults. Numbers of
captures at burrows were best correlated with large numbers
of herbaceous species and high stem counts of Pinus palus-
tris, Ouercus laevis, and Diospyros virqiniana: there was
little correlation with ground cover characteristics.
Immediate responses of Podomys to prescribed burns were not
spectacular, but in the long term, populations appeared to
be more stable on sites that were burned periodically than
on areas not burned. Implications for conservation of
Podomys on the Ordway Preserve also were discussed.
Xll

INTRODUCTION
An understanding of home range, diet, burrow use, and
other interactions between an animal and its physical en¬
vironment is crucial for the management and conservation of
threatened and endangered species. However, little is known
about the ecology of Florida's only endemic mammal, the
Florida or gopher mouse. This mouse, Podomys floridanus
(Rodentia: Muridae), is listed as Threatened by the Florida
Committee on Rare and Endangered Plants and Animals (Layne,
1978) and as a Species of Special Concern by the Florida
Game and Fresh Water Fish Commission (1987). Podomys flori¬
danus lives in a narrow range of habitats; my study, and
those by Layne at the Archbold Biological Station in High¬
lands County, suggest that P. floridanus populations are
higher in the southern sand pine scrub than in the high pine
lands (also known as turkey oak-longleaf pine sandhills or
high pine hills). The special listing of P. floridanus is
due to destruction of high pine and scrub habitats and to
the decline of gopher tortoise (Gopherus Polyphemus) popula¬
tions. Nash (1895) stated that high pine was the most
extensive of the five habitats he recognized in Florida; by
1

2
1982 approximately 57% of this habitat had been lost (Auf-
fenberg and Franz, 1982).
In the high pine lands of northern Florida, the Florida
mouse is associated closely with burrows of the gopher
tortoise (Blair and Kilby, 1936; Humphrey et al., 1985;
Brand, 1987). My preliminary field studies showed that
populations of Florida mice were relatively high and stable
on certain sandhills, whereas populations fluctuated to a
greater degree on other sites. I was intrigued by the
differences I observed in these populations. How did these
sandhills differ from each other? Why were certain tortoise
burrows favored nesting sites, year after year, whereas
other burrows never were used? In an attempt to determine
microhabitat preferences in the high pine habitats, I tried
five approaches:
1) Do Podomvs in the high pine depend exclusively on
tortoise burrows? Mouse-use of tortoise burrows
was monitored over a five-year period so that
long-term "burrow histories" could be constructed.
2) How large are home ranges in the high pine? Home
range areas of Podomvs in high pine were measured.
Do dietary data explain why some tortoise burrows
are preferred over others? Diets of these mice
were observed in the field and laboratory. A
supplemental feeding experiment was performed to
test the hypothesis that population size is food-
limited (i.e., does increased food supply produce
an increase in numbers and body size of mice and
an increase in the length of the breeding sea¬
son? ) .
3)

3
4) Is the local distribution of these mice predic¬
table? Physical parameters of burrows, and the
vegetation surrounding them, were quantified.
5) Do populations respond positively to burns?
Effects of prescribed burns on populations of
Podomvs were determined.

TAXONOMY AND SYSTEMATICS
Podomys floridanus is the only mammalian species en¬
demic to Florida. The species was described as Hesperomvs
floridanus by Frank Chapman in 1889, with the type locality
in Gainesville, Alachua County.
Podomys floridanus has long been recognized as distinct
from North American species of Peromyscus. Osgood (1909)
considered Podomys a distinct subgenus within the genus
Peromyscus. Apparently Hooper (1958) was the first to
suggest that the closest relatives of Podomys were not the
cotton mouse and other local Peromyscus. but peromyscines in
Central America. In Carleton's (1980) major revision of the
genus Peromyscus. Podomys was raised from subgenus to genus.
Carleton suggested that Podomys shared a common ancestor
with Habromys and Neotomodon from Mexico and Guatemala;
these three genera grouped together consistently in his
cladistic analysis of 79 morphological characters.
The proposed affinity between Podomys and mice from
Central America is supported by three lines of evidence:
1) Reproductive tract. The baculum of Podomys is
relatively small and slender, compared to that of
northern Peromyscus. Linzey and Layne (1969)
examined additional features of the male reproduc¬
tive tract and noted reduced vesicular and
4

5
prostate glands, enlarged ampullary glands, and
other features that suggested affinity with Habro-
mvs.
2) Ectoparasites. The "typical and only common flea"
of Podomys, described by Johnson and Layne (1961),
is Polyqenis floridanus. which most closely resem¬
bles a species found in South America. According
to Wenzel and Tipton (1966: 718), "Polyqenis is an
"expanding" South American genus which has, quite
clearly, dispersed into Middle America along with
complex-penis-type Cricetinae and caviomorph
rodents from South America".
3) Electrophoresis. In their analysis of genic
variation among Peromvscus. Smith et al. (1973)
considered Podomys one of the least variable
species of the group. One possible explanation
for this low variability is the maintenance of low
numbers in a relictual population.
The fossil record of the Florida mouse is limited to
six Rancholabrean assemblages in this state, so there is not
much light shed on the distribution of Podomys prior to the
Illinoisan. However, a western affinity might be indicated
by a Pleistocene species (Peromvscus oklahomensis) from the
Illinoisan and Sangamonian of Oklahoma and Texas, that might
be related to Podomys (Stephens, 1960). Unfortunately, only
a few isolated teeth of this extinct species have been
found.
Podomys is not unique among the flora and fauna of
Florida in its apparent affinity to Central America. Several
plants (such as Opuntia. the prickly pear) and several
reptiles, birds, and mammals are considered species of
western affinity. Some, such as Gopherus polyphemus. show a
disjunct distribution (Blair, 1958). Among the mammals, Webb
and Wilkins (1984) found that 15-28% of the mammals from

6
five Pleistocene sites in Florida were neotropical species.
They suggested that the arrival of most of these species
could be explained by a Gulf Coastal Corridor, which served
as a pathway for southern and western immigrants in the
early Pleistocene.

DISTRIBUTION
In recent times P. floridanus occurred from St. Johns,
Clay, Alachua, Putnam, Suwannee, and Taylor counties south
to Sarasota, Highlands, and Dade counties, with isolated
populations in Franklin County and on Merritt Island (Os¬
good, 1909; Layne, 1978; Stout et al., 1979). A survey of
the present distribution of Podomvs is urgently needed.
Podomvs is the characteristic mouse of some of the most
xeric habitats in Florida. It is most common in sand pine
scrub and high pine, although it also is found in slash
pine-turkey oak and coastal strand associations (Bangs,
1898; Layne, 1978; U.S. Dept. Agrie., 1980). On Fig. 1, I
have mapped the distribution of scrub and high pine, as
indicated by the Soil Conservation Service (U.S. Dept.
Agrie., 1980), and localities from which Podomvs have been
reported (Layne, 1978). Clearly there is a close associa¬
tion between Podomvs and these two habitats. Layne (in
press) has suggested that the scrub was the ancestral home
of this mouse, with the high pine a secondary habitat.
In some areas, P. floridanus and P. aossypinus appear
to coexist (Layne, 1970). In Putnam County, however, trapp¬
ing by Suzanne Brand (1987), John Eisenberg (1983; 1988),
7

8
Distribution of Podomys floridanus (triangle
designates type locality). Stippling represents
distribution of high pine and scrub habitats.
Fig. 1.

9
Eric Milstrey (1987), and me has indicated a clear separa¬
tion between these two species, with P. qossypinus occurring
only in lower, wetter microhabitats, and Podomvs only in
sandhills, or along edges between sandhills and old pastures
or sandhills and xeric hammocks.

MATERIALS AND METHODS
I conducted this research on the Katharine Ordway
Preserve-Carl S. Swisher Memorial Sanctuary, located about
4 km E of Melrose in Putnam County. The preserve, 37 km2 in
size, is administered jointly by the Florida Museum of Na¬
tural History, the University of Florida's School of Forest
Resources and Conservation, and the Nature Conservancy.
Approximately one-third of the Ordway Preserve is
covered by high pine land (sensu Nash, 1895). This habitat
is characterized by an open canopy of longleaf pine (Pinus
palustris) and turkey oak (Ouercus laevis), although other
trees, including sand pine (P. clausa). Chapman's oak (Q.
chapmanii). and bluejack (Q. incana), also are present. The
understory consists of scattered shrubs, and herbs and
grasses. On Ordway the characteristic grass of the high
pine land is wiregrass (Aristida stricta); other common
grasses include pineywoods dropseed (Sporobolus iunceus),
six species of bluestem (Andropogon spp.), and three species
of threeawn (Aristida spp.). Additional grasses identified
on the preserve are included in the list compiled by David
W. Hall and Richard Franz (unpubl. data). Reindeer moss
10

11
(Cladonia and Cladina spp.) is common. The herbs I
identified on my study sites are listed in the Appendix.
The soils of high pine are sandy and well-drained.
There are many patches of bare ground, which become covered
with oak leaves, pine needles, tree branches, and other
debris in the absence of fire. Under the pines, the needle
litter is several inches deep; in areas where the turkey
oaks predominate, leaf litter is distributed unevenly by the
wind. Burrows of the gopher tortoise (Gopherus polyphemus)
are prominent features on this landscape, although popula¬
tion densities probably were higher in coastal areas before
human development (Carr, 1940; Auffenberg and Franz, 1982;
Cox et al., 1987).
Other mammals found on the high pine land are the
opossum lDidelphis virginianus), pocket gopher (Geomys
pinetis), fox squirrel (Sciurus niger), flying squirrel
(Glaucomvs volans1, armadillo (Dasypus nóvemeinctus), rac¬
coon (Procyon lptor), and white-tailed deer (Odocoileus
virginianus). Eisenberg (1988) and Franz (1989) compiled
lists of mammals known from the various habitats of Ordway.
Carr (1940) listed the amphibians and reptiles of high pine
habitats. On my study sites, I saw gopher frogs (Rana
areplata), oak toads (Bufo guercicus), fence lizards (Scelo-
porus undulatus), six-lined racerunners (Cnemidophorus
sexlineatus), pine snakes (Pituophis melanoleucus), crowned
snakes (Tantilla spp.), southern black racers (Coluber

12
constrictor^, coachwhips (Masticophis flagellum^, coral
snakes (Micrurus fulvius), pigmy rattlesnakes lSistrurus
miliarius1, eastern diamondbacks (Crotalus adamanteus), and
tortoises (G. polyphemus). Birds common to high pine were
listed by Nelson (1952), who thought the avifauna of the
high pine did not differ substantially from those of oak
hammocks and other nearby communities. However, Maehr et
al. (1982) suggested that the high pine might support a
greater density of birds than flatwoods and xeric hammocks.
I did not attempt to identify birds on my study sites in
detail.
On the Ordway Preserve, all locations at which P.
floridanus have been found are in high pine habitats, or at
the ecotone between high pine and hammock or high pine and
old pasture (Brand, 1987; Eisenberg, 1988) (Fig. 2). I
trapped mice on four independent high-pine sites, designated
Anderson-Cue (AC), Blue Pond (BP), Longleaf Pine Pasture
(LP), and Smith Lake (SL). The history of fire on these
sites is described below in the Prescribed Burns section.
Smith Lake is my longterm study site; Eisenberg and Franz
started research projects there in 1983, marking tortoise
burrows and trapping Florida mice, gopher frogs, and other
burrow inhabitants. I trapped mice at SL from 1984 through
1988. SL is approximately 16.8 ha in size. Vegetation
consists of P. palustris. Q. laeyis, Q. chapmanii, Q. qemin-
ata, and grasses and forbs characteristic of the high pine.

Distribution of Podomvs on the Ordway/Swisher Preserve. Locations of
Anderson-Cue (AC), Blue Pond (BP), Longleaf Pine (LP), and Smith Lake
(SL) are shown. Dots represent additional locations where Podomys
have been captured.
Fig. 2.


15
The Anderson-Cue sandhill is the highest in elevation of the
four sites; typical vegetation there includes P. palustris
and Q. laevis, with fewer Q. qeminata than on SL. I worked
there from 1986 to 1988. AC was burned in the winters of
1983, 1986, and 1989. It was not included in my study of
prescribed burns because I did no pre-burn trapping there.
The Longleaf Pine Pasture Sandhill is south of the pine
pasture surrounding the old Swisher Estate (Fig. 2). Part
of this sandhill, the most disturbed of my study areas, has
been planted in slash pine. Dominant plants here include
maidencane (Panicum hemitpmon) and centipede grass (Eremoch-
loa ophiuroides), which are not typical of natural high pine
vegetation. There are many turkey oaks and several large
patches of gopher apple, but fewer longleaf pines, pawpaws,
and other high pine species. I worked on LP for only one
trapping season for the purpose of gathering additional
information regarding prescribed burns. The fourth study
site, Blue Pond, was logged intensively. There are few
longleaf pines, and turkey oaks are by far the most numerous
trees. Ground cover is sparse in some areas but rather
dense in others. Species composition of the ground cover
appeared similar to that of SL.
As described in the following chapters, I trapped P.
floridanus at tortoise burrows and on grids using standard
mark-and-recapture techniques (Davis, 1956). While trapping
at burrows, I numbered, marked, and mapped each burrow, and

16
then placed a pair of Sherman traps (baited with oatmeal) at
the entrance, or just inside the mouth, of the burrow.
Animals were marked by toe-clipping and then weighed. I
also noted sex, reproductive condition, and presence of
ectoparasites. Animals were released near the burrow en¬
trance, and in most cases I recorded whether they entered
the burrow or ran elsewhere. I collected the same data on
grids. I placed one grid (designated SL) on my study site
on the south side of SL; two more grids (ACI and ACII) were
set north and east of the Anderson-Cue ponds. Grids were
more than 100 ra apart to reduce movement of animals between
grids. Each grid consisted of 10 columns and 10 rows 10 m
apart (area = 10000 m2), with a single Sherman at each
intersection. Each tortoise burrow and grid location was
given a unique number and marked with forestry flags. On
the burrow flags I also attached an aluminum tag (Al Tag,
Forestry Suppliers, 205 W. Rankin Street, Jackson, Missis¬
sippi 39204) bearing the burrow number. These tags survived
all but the hottest burns, thus serving as more permanent
markers of burrow locations.
Traps were opened near sundown and checked at or short¬
ly following sunrise. Because of temperature extremes,
morning condensation, and (in some months) heavy rain, I
found it necessary to check traps in the morning as quickly
as possible. Animals developed hypothermia when ambient
temperatures reached 10 C; I extended the trapping season

17
into November by insulating traps with a handful of excel¬
sior, wood shavings used in packing glassware. Mice com¬
press the excelsior into a pad on top of the treadle.
Spanish moss also can be used, but I found the wood to be
superior because it does not absorb moisture. I did not
trap in December or January; fieldwork resumed when warm
weather resumed (February-April) .
I measured trapping effort as trapnights (one trapnight
being one trap/night). Results of the vegetation analyses
were run on SYSTAT at the IFAS Consulting Center at the
University of Florida; other statistics were performed on a
hand-held calculator. Statistical methods are described in
each chapter.

USE OF TORTOISE BURROWS
Dependence on Burrows
Previous students of peromyscine behavior and morphol¬
ogy concluded that Podomvs was primarily a terrestrial
species; although capable of climbing and digging, Podomvs
climbed and dug less than several species of Peromvscus to
which it was compared (King, 1968; Layne 1970; Layne and
Ehrhart, 1970; Siegel and Van Meter, 1973). In some areas,
these mice seemed capable of excavating their own burrows;
Lee (1968) stated that P. floridanus built burrows greater
than 120 cm in depth in scrub habitats, and Starner (1956)
trapped one animal in a small burrow that she thought might
have been dug by the mouse (in the ecotone between the mesic
hammock and longleaf pine flatwoods at San Felasco Hammock,
Alachua County). Layne and Ehrhart (1970) suggested that
Podomvs rarely dug its own burrows, but Layne (in press)
noted high populations of Florida mice in scrubs where
tortoises were absent or occurred in low numbers. However,
observations by Eisenberg (1983), Brand (1987), and myself
suggested a much closer relationship between Podomvs and
Gopherus in the high pine in Putnam County.
18

19
Blair and Kilby (1936) were the first to note in the
literature that Podomvs floridanus used burrows of Gopherus
Polyphemus in northern and central Florida. They saw mice
using active burrows (i.e., burrows that had tortoises in
them) in an old field in Alachua County. Three of the 15
mice captured by Starner (1956) were taken from traps placed
inside entrances of tortoise burrows. Milstrey (1987:115)
reported that "Florida mice were continuously present" in
many tortoise burrows at his study site in Putnam County.
Johnson and Layne (1961) also noted that Podomvs inhabited
tortoise burrows, and possibly burrows of pocket gophers
(Geomvs pinetis). At other sites Podomvs used burrows
constructed by Peromvscus polionotus, Sicrmodon hispidus, G.
pinetis. and Dasypus nóvemeinctus (Gentry and Smith, 1968;
Layne, in press). However, in my 33,385 trapnights on
burrows and grids at Ordway I have observed Florida mice
entering small holes of unknown origin only three times
(twice on SL and once on AC), and I have no evidence that
they inhabited logs or other shelters.
I found that mice almost always ran down tortoise
burrows when released. For example, I tallied escape re¬
sponses of 18 Podomvs caught on the AC grid (and therefore
released at grid coordinates, not at burrows). Out of 35
captures, mice ran down tortoise burrows 25 times; they hid
in the base of turkey oaks or in fallen logs and other
debris 10 times. In my regular trapping sessions, I never

20
saw mice climb vegetation as an escape response; Eisenberg
(pers. comm.) noted one such response in 1989. However, my
field observations have confirmed climbing ability by these
mice (Jones, 1989, and Home Range below). In T-maze tests
of arboreality performed by C. A. Langtimm at SL (in which a
T-maze was propped against an oak), most mice jumped off the
T, but the animals who chose to climb oaks reached heights
of 8 m (C. A. Langtimm, unpubl. data). Clearly these ani¬
mals are capable of climbing, even though the time spent on
this activity in any one night remains unknown.
In preliminary studies on the Ordway Preserve, Eisen¬
berg (1983) reported that trapping success was much higher
if traps were placed at the mouths of tortoise burrows (33%)
than on transects (0.4%). Humphrey et al. (1985) caught 113
mice in 1,144 trapnights at 51 burrows, also at Ordway. In
146 trapnights on transects—in which traps were not placed
at burrows--I caught no mice. At the SL burrows in August,
September, October, and November 1983, Eisenberg (unpubl.
data) had trapping successes of 35, 43, 62, and 44%, respec¬
tively. I calculated trapping success using adjusted trap¬
nights, which indicated how many traps were available after
subtracting half the number of traps sprung and containing
non-target species from the total number of traps set. My
trap success for Podomvs ranged from 0 to 16%, with SL being
the most successful (2-16% with a mean of 7%) (Fig. 3).
Variations in total trapnights (within and among sites)

Fig. 3. Trapping success (%) and minimum numbers of individuals (MNI) (adults
and subadults) of Podomys at burrows on the Smith Lake Sandhill.
Monthly trapping effort was unequal among months.

TRAPPING SUCCESS (%) AND MNI
MONTH
to
to

23
reflect my attempt to maximize sample size by trapping at
all tortoise burrows on each sandhill until burrows disap¬
peared.
On all four sandhills, traps at burrows occasionally
were sprung by falling branches or by insects removing bait.
A large number of sprung traps was an occasional problem at
AC, but a more major one at BP, particularly on the north
side. There I identified tracks of armadillos, raccoons,
and dogs, but I suspected dogs were the major culprits. One
morning I found two traps covered with urine and several
times I saw dogs on the sandhill, which was the closest of
my sites to the periphery of the preserve. It is also worth
noting that I found more carcasses of turtles and tortoises
here than on any other study site. Following a suggestion
of M. E. Sunquist, I fashioned "croquet hoops" out of alu¬
minum welding rods and placed them around the traps. They
helped decrease the number of traps moved, but did not
always prevent closing of trap doors.
In traps at burrows, the most frequently-captured non¬
target species was Rana areolata (Table 1). At SL, from
September 1984 to the end of the study, gopher frogs were
captured 105 times. Other occasional captures were camel
crickets (Ceuthophilus sp.), centipedes, southern toads
(Bufo terrestris. once on AC, LP, and SL), six-lined race-
runners (Cnemidophorus sexlineatus ^, and a young rabbit
(presumably Svlvilaqus floridanus. on SL). All of these

24
TABLE 1. Numbers of captures of non-target species in traps
at burrows (22,085 trapnights) and on grids (11,300 trap-
nights). Captures of insects and centipedes were not
counted. Scientific names are listed in text.
Burrows
Grids
camel crickets
grasshoppers
1 spider
cockroaches
3 southern toad
centipedes
129 gopher frog
3
six-lined
racerunner
3 six-lined racerunner
2
fence lizard
1 cotton mouse
1
black racer
38 flying squirrel
2
rice rat
1 rabbit
3
cotton rat
148 flying squirrel

25
species are known to use Gopherus burrows (Jackson and
Milstrey, 1989). In 1988 I began catching flying squirrels
(Glaucomvs yolans) at the AC and SL burrows; at SL I cap¬
tured squirrels 27 times from May to October, 15 times in
May alone. These results must indicate some change in the
Glaucomvs population that resulted in additional terrestrial
foraging, or perhaps they reflect learned behavior of ani¬
mals who found oats at traps on grids. At any rate, I
observed only one flying squirrel enter a burrow upon re¬
lease, and I do not believe they reside in tortoise burrows
on these sandhills.
At study sites other than Ordway, additional murids,
including P. qossypinus and P. polionotus. visit tortoise
burrows (Jackson and Milstrey, 1989). In 22,085 trapnights
at burrows on the AC,BP, LP, and SL sandhills I caught only
one cotton mouse (P. qossypinus. on AC). In other Ordway
studies, Eisenberg (1983) and Brand (1987) found Podomys in
the high pine and Peromvscus in the hammocks and flatwoods.
Clearly, Podomys is the only rodent that regularly inhabits
tortoise burrows in these sandhills. At burrows in other
Ordway habitats, the absence or lower numbers of Podomys
might represent a case of competitive exclusion; after the
1987 drought Eisenberg (pers. comm.) reported trapping
Florida mice in hammocks closer to lakes than in previous
years, and Peromvscus were absent.

26
For comparative purposes, trapping results on two grids
on AC and one grid on SL also are shown (Fig. 4). Trapping
success on grids ranged from 0 to 7% (AC) and from 0 to 6%
(SL); mean trapping successes were 0.47%, 0.70%, and 3.6%
for ACI, ACII, and SL. Traps on grids were more likely to
be sprung by rain, falling branches, pine cones, and rac¬
coons than were traps at burrows. Only one Florida mouse
(female 18, a juvenile known to be alive 36 days at SL) was
found on a grid and not during burrow trapping. Flying
squirrels were caught more frequently (Table 1), usually at
traps near the bases of pines, oaks, or dead trees. One
cotton rat (Siomodon hispidus) was captured once at SL; this
was a juvenile male (which I marked) presumably dispersing
across the high pine. The other cotton rats and the rice
rats (Orvzomvs palustris) were caught on the AC grids on 28
and 29 June 1988; their appearance on the grids might re¬
flect drought-related dispersal out of the vegetation sur¬
rounding the AC ponds. Centipedes occasionally were caught
on all grids. On ACI I also captured grasshoppers, cock¬
roaches, fence lizards (Sceloporus undulatus), six-lined
racerunners (C. sexlineatus), and a black racer (Coluber
constrictor).
Description of Burrows
The size and configuration of tortoise burrows have
been described by Hallinan (1923) and Hansen (1964). Blair

27
en
en
ni
o
o
D
en
O
z
CL
CL
<
ce
I-
1987 1988
MONTH
en
en
LU
o
o
Z
en
o
z
o.
CL
<
ce
1 987
MONTH
1 988
Fig
4.
Trapping success (expressed as the total number of
captures/adjusted trapnights) for Podomys on the
AC (top) and SL (bottom) grids.

28
and Kilby (1936) noted the use of a small hole approximately
182 cm from the entrance of a burrow. However, the only
description of structures used by Podomvs was that of Layne
(in press), who described nests, nest chambers, and small
tunnels found in two tortoise burrows in Alachua County.
Richard Franz and I excavated five burrows on Ordway in
order to further describe structures utilized by mice.
Typically, mice used small U-shaped tunnels constructed off
one side of the tortoise burrow, with both entrances to the
U opening into the main burrow. We also noted use of tun¬
nels we dubbed "chimneys", which ran from the ceiling or
side of the burrow to an outside opening a meter or two past
the main entrance. In all five burrows, mouse activity
appeared concentrated in the upper 2 m of the burrow.
Additional details of the actual excavations were discussed
by Jones and Franz (in prep.). Chimneys allowed occupation
of burrows after the main entrance has collapsed, and might
serve as escape routes from predators. Although not all
burrows have chimneys (at my four study sites from 6.8% (AC)
to 26.9% (BP) of the burrows had chimneys), the presence of
chimneys is an indicator of past or present occupation of
tortoise burrows by Podomvs. Exploratory trapping for mice
in the sandhills should not omit observation of these small
holes.
To determine the pattern of dispersion of tortoise
burrows on the SL sandhill, I used methods described by

29
Brower and Zar (1984). In this procedure, a grid is placed
over a map of the study area and numbers of squares contain¬
ing no burrows, squares containing one burrow, and etc., are
counted. I used a grid with squares 35 mm by 35 mm (which
corresponds to 1225 m2 on the map) . My SL study site con¬
sisted of 93 squares with a mean of 1.46 burrows per square
(or 0.11 burrow/m2). I included all 136 burrows monitored
from 1983 through 1988, without consideration of burrow
classification or tortoise demography; that is, I considered
only burrow locations without burrow size, duration of
residence, or other characteristics. The data fit the
Poisson distribution, leading to the conclusion that the 136
burrows on SL are randomly distributed (Table 2). Another
method to determine whether distribution is random is the X2
test, for which I obtained a value of 0.547 (not signifi¬
cantly different from random at p = 0.05). As a third me¬
thod, I tried the Morisita's index of dispersion, which is
calculated as I = n [ (2X2-N) / (N(N-l) ) ] where n is the number
of squares, N is the total number of burrows counted on n
squares, and EX2 is the squares of the numbers of burrows
per plots for all plots. A value of 1.0 for I indicates
random dispersion; value of 0 indicates uniform dispersion.
I obtained I = 0.95, which indicates random dispersion.
Finally, I calculated the ratio of observed to expected
density; I obtained a value of 0.94, which is close to the
value of 1.00 which again indicates random distribution

30
TABLE 2. Data describing distribution of burrows on SL
(top) and AC (bottom) with the probabilities expected from
the Poisson distribution (with means of 1.5 and 1.7 burrows
per square). X indicates number of burrows per square; f(X)
indicates the observed frequency; p(X) is the observed
probability; and P(X) the expected Poisson probability.
X
f(X)
P(X)
em
0
20
0.215
0.223
1
33
0.354
0.335
2
24
0.258
0.251
3
10
0.108
0.126
4
5
0.054
0.047
5
1
0.011
0.014
0
15
0.259
0.183
1
13
0.224
0.311
2
13
0.224
0.264
3
12
0.207
0.150
4
3
0.052
0.064
5
0
0
0.022
6
0
0
0.006
7
2
0.035
0.001

31
(Brower and Zar, 1984). The 101 AC burrows also were dis¬
persed randomly (X2 = 5, significant at p = 0.10; I = 0.98;
ratio of observed to expected = 1.29).
Observations of dispersion of other gopher tortoise
burrows vary. McRae et al. (1981) and Auffenberg and Franz
(1982) recognized burrows clumped as colonies, whereas
others reported random distributions (e.g., Kushlan, 1982;
Kushlan and Mazzotti, 1984). Whether these discrepancies
reflect variations in tortoise behavior or in data collec¬
tion remains unknown.
Podomvs and the Burrow Community
In the high pine, Gopherus burrows provide shelter from
the fires and extreme fluctuations in temperature and rain¬
fall that characterize this environment. Burrows of the
southeastern pocket gopher (Geomys pinetis1 also are common
in the high pine, but the two types of burrows are used by
different species (Funderburg and Lee, 1968). Eisenberg
(1983) suggested that G. polyphemus could be considered a
keystone species in the high pine due to the numerous and
diverse species that use its burrows. Franz (1986) listed
19 species of amphibians and reptiles that used the SL bur¬
rows. The most recent state-wide list of species associated
with Gopherus burrows consists of 60 vertebrates (Amphibia,
Reptilia, Aves, and Mammalia) and 302 invertebrate species

32
(Gastropoda, Malacostraca, Arachnida, Chilopoda, Diplopoda,
and Insecta) (Jackson and Milstrey, 1989).
Besides using burrows as shelter, I suspect Podomvs
takes advantage of the invertebrate commensals and visitors.
My captive mice readily accepted live crickets and meal¬
worms. Milstrey (1987) described Podomvs eating blood and
body fluids of engorged ticks (Ornithodoros turicata) that
parasitize Gopherus and Rana areolata.
Florida mice are presumably preyed upon by birds,
foxes, bobcats, and raccoons (Layne, 1978; Maehr and Brady,
1986; Wassmer et al., 1988). I suspect that in the high
pine snakes are major predators, although predation by
snakes has not been documented. I occasionally saw coral
snakes (M. fulvius) near burrows, but omitted the species
from consideration because coral snakes do not prey upon
mammals (Jackson and Franz, 1981). Mushinsky (1987) sum¬
marized predation records for snakes; some of the species
known to take mammals include Coluber constrictor. Drvmar-
chon coráis. Elaphe obsoleta, and Pituophis melanoleucus.
Several species of snakes utilize tortoise burrows at Ordway
and must be considered potential predators of Podomvs,
including C. constrictor. E. guttata, P. melanoleucus,
Masticophis flagellum. D. coráis. Sistrurus miliarius, and
Crotalus adamanteus (Franz, 1986; Timmerman, 1989). In
particular, eastern diamondbacks are known to take Peromvs-
cus, and Timmerman (1989) considered Podomvs a potential

33
prey item. During my time at Ordway, I saw only one dia-
mondback in the high pine (an adult female near BP burrow
#319, 6 November 1988). Timmerman (1989) examined habitats
used by eight eastern diamondbacks at Ordway from 1985 to
1989. His snakes used high pine and old field habitats less
than expected (approximately 13% of all observations) al¬
though these two habitats comprised more than 60% of the
area available. The diamondbacks spent little time under¬
ground, and only 16% of the burrow locations were in Gopher-
us burrows. However, as Timmerman pointed out, 75% of the
locations were in dens of the armadillo, which frequently
takes over old Gopherus burrows. Tortoise burrows are used
as winter refugia and, in the fall, as parturition sites,
leading to the suggestion that Podomvs might be important in
the diet of young diamondbacks (Timmerman, 1989).
Mortality rates of Podomvs due to disease and parasites
are unknown. Thirty-two species of endo- and exoparasites
of Podomvs have been reported (Layne, 1963, 1968a; Lichten-
fels, 1970; Whitaker, 1968). Milstrey (1984, 1987) sum¬
marized recent studies of the soft tick Ornithodorus. a
known carrier of relapsing fever that feeds on residents of
gopher burrows, but, as he points out, the relationships
between the vertebrates and invertebrates of these burrow
systems are still poorly understood.

34
Life History
I examined demographic data for my largest sample, the
255 P. floridanus marked at the SL burrows, August 1983 to
November 1988. Of this total, 52 females and 44 males were
marked in 1983. Eighty-one (38 females, 43 males)of the 255
(32%) were captured only once. I calculated minimum trap-
pability ((number of captures - 2)/(possible captures -2))
for animals captured three or more times (Hilborn et al.,
1976) .
Individual trappability ranged from 14 to 100%. Aver¬
age trappability was 58% for 27 mice in 1983, 55% for n = 17
in 1984, 58% for n = 16 in 1985, 62% for n = 16 in 1986, 48%
for n = 13 in 1987, and 60% for n = 24 in 1988. The average
for all five years was 57%. According to Hilborn et al.
(1976), when trappability = < 50%, estimates of minimum num¬
bers of individuals (MNI) by direct enumeration will under¬
estimate actual population size; estimates become more
reliable as trappability increases. Trappability on grids
was lower than on burrows. For example, individual trap¬
pability on the SL grid ranges from 38 to 50%.
The reproduction, growth, and development of Podomvs
were described by Layne (1966). He demonstrated that sea¬
sonal distribution of pregnancies in a wild population in
Alachua County was bimodal, with the major peak occurring
between August and November. At SL, I classified young
animals as juveniles if the gray and white juvenile pelage

35
was still present, and as subadults if the dorsum was gray
with gold and agouti hairs appearing along sides and flanks
(minimum weights of subadults were approximately 25 g).
Numbers of young marked (Fig. 5) and numbers of reproductive
adults (Fig. 6) at SL indicate that seasonality of reproduc¬
tion at Ordway also peaked in the late summer and fall,
although juveniles and reproductive adults might appear in
small numbers at other times of the year (Layne, 1966).
Juveniles usually were first captured at the mouths of
burrows at approximately three weeks of age. At this point
the young were still completely gray and white and weighed
approximately 9 to 14 g (Layne, 1966). In some cases young
appeared at burrows with females (and sometimes males) who
were long-term residents, but I had no way of determining
parentage of some juveniles and almost all subadults. This,
and the fact that many juveniles disappeared before comple¬
tion of adult molt, suggests that both pre- and post-
reproductive dispersal occur.
Captive animals breed year-round (Dice, 1954; Layne,
1966; Drickamer and Vestal, 1973). In captivity mean litter
size (1.7-2.6) is slightly smaller than that reported (3.1)
for wild-conceived litters (Dice, 1954; Layne, 1966; Rood,
1966; Drickamer and Vestal, 1973; Glazier, 1985). Maternal
care in captivity consisted of creating nests, plugging
entrances to nest boxes, nursing and grooming young, and
herding juveniles into the nest, as described previously by

36
MONTH
Numbers of juveniles (dots) and subadults
(circles) marked on the SL sandhill, expressed as
minimum numbers of individuals/number of
trapnights (top) and as numbers of individuals
(bottom).
Fig. 5.

% PREGNANT
37
MONTH
—*□— 1984
MONTH
1984
1985
1986
1987
1988
Reproductive data for Podomys trapped on the SL
sandhill, 1983-1988: the proportion of adult
females who were pregnant (top) and the proportion
of males who were scrotal (bottom) are shown.
Fig. 6.

38
Layne (1966, 1968b). In my captive colony 8 females and 5
males produced 54 young in 24 litters; 11 litters were born
to a single pair. Litter size ranged from 1 to 4 (mean =
2.25). The distribution of births was: 8 in September; 4 in
July; 3 in October; 2 in March, August, and December; 1 in
January, February, and November. Four neonates have not
been sexed at this time; the remaining young consisted of 24
females, 19 males, and 7 infants who died before gender was
recognized.
Longevity of Podomvs in the wild is unknown. Keim and
Stout (1987) reported that a male captured at approximately
three weeks of age lived in captivity for 7 years and 4
months. At SL, 12 females and 10 males (8.6% of the animals
marked) were present for more than 360 days (Table 3).
Seven females and one male were first marked as juveniles;
20 individuals were first found on the south side of SL and
two were marked on the north and later migrated south. In
captivity, two- and three-year-olds are continuing to repro¬
duce .

39
TABLE 3. Longevity records exceeding 360 days for Podomys
floridanus at SL. ID is the identification number, YR =
year marked, A = age when marked (Juvenile, Subadult, or
Adult), and P = persistence (in days) as indicated by both
trapping methods. Asterisk indicates trap mortality.
Females
Males
ID
YR
A
P
ID
YR
A
P
43
83
S
445
39
83
A
905
55
83
J
409
48
83
A
570
65
83
J
649
63
83
S
410
23
84
A
467
11
84
A
920
30
84
A
403
32
84
A
620
61
86
J
360
98
85
A
401*
10
87
J
588
80
86
S
515
19
87
J
385
98B
86
S
500
26
87
J
554
42
87
A
457
47
87
A
547
54
87
J
412
58
87
A
392
64
87
J
504

HOME RANGE
The concept of home range most widely accepted by
mammalogists is that defined by Burt (1943:351) as "that
area traversed by the individual in its normal activities of
food gathering, mating, and caring for young. Occasional
sallies outside the area . . . should not be considered as
in part of the home range. The home range need not cover
the same area during the life of the individual." The
concept of territoriality, on the other hand, involves
domination by either overt or covert social interaction. As
defined by Kaufmann (1983:9), a territory is "a fixed por¬
tion of an individual's or group's range in which it has
priority of access to one or more critical resources over
others which have priority elsewhere or at another time."
Among peromyscines, there is considerable variation in
descriptions of both home range sizes and degrees of ter¬
ritoriality (Eisenberg, 1968; Stickel, 1968). For P. flori-
danus, home range and territoriality have not been des¬
cribed. I examined the size of home ranges on the Ordway
sandhills with three methods: fluorescent powder, trapping
on grids, and trapping at burrows.
40

41
Fluorescent Powder
The use of fluorescent powder to track small rodents
and to determine movement patterns was introduced in the
sandhills of Nebraska (Lemen and Freeman, 1985, 1986). I
applied this technique to examine size of nightly activity
areas of Podomvs on the SL sandhill.
I powdered five animals on 25 May 1985, three on 14
July 1985, and three on 15 June 1987. The first eight ani¬
mals were trapped at burrows and held until 2100 EST the
following evening; the last three were removed from traps on
the grid at 0230 EST, powdered, and released immediately.
Each animal was placed into a plastic ziplock bag with a
fine, nontoxic fluorescent powder (green, pink, or yellow),
gently shaken until completely covered, and released at the
point of capture. At a later time, I returned with the
fluorescent light and marked the powder trails with forestry
flags. Trails then were mapped using a tape measure and
compass.
The mice showed no ill effects from this technique by
loss in weight or other signs of poor condition. One female
was so little traumatized by the procedure that she caught
and consumed a moth immediately after her release. However,
with the exception of the moth incident, I did not obtain
additional information regarding feeding.
Powdering was of some use in that it showed animals
utilizing fallen trees and branches as runways, as I had

42
observed in escape responses during trapping. Douglass and
Reinert (1982) showed that utilization of fallen logs by
Peromvscus leucopus was not an artifact of trapping but part
of the animals' normal movements. Additional evidence of
climbing was indicated by the powder trail of an adult male
(#39), which extended 36 cm up a turkey oak snag.
The powder indicated only one case of social interac¬
tion. On 15 June 1987 females 10 and 47 were caught in the
same trap. Female 47 was powdered, released, and recaptured
later that morning. Again, she was found in the same trap
with #10, who also had become covered with powder. Female
47 was first marked as an adult on the SL grid on 19 April;
she remained at the grid and surrounding burrows until at
least October 1988. Number 10 was a younger female marked
as a juvenile on 12 April at a burrow located on the grid;
by September she had moved to an area west of the grid.
Possibly #10 was a daughter of #47. Use of radioisotopes,
DNA fingerprinting, and other techniques would be more
appropriate and efficient methods to examine family rela¬
tionships and dispersal of young Podomvs.
Distances traveled by mice, as indicated by powder
trails, are shown in Table 4. Trails for two animals were
very short (< 10 m), but I believed that a single night's
excursion exceeded 10 m. By trapping on 14 July 1985 and 15
June 1987, I discovered powdered mice at locations to which
no powder trails led. Thus, the powder trail showed only a

43
TABLE 4. Activity data from ten Podomvs dusted with fluo-
rescent powder at SL. ID indicates sex and identification
number of mouse; R indicates the burrow or grid station at
which an animal was caught and released; ORI indicates
initial direction from release point (in degrees); DIST is
the greatest distance (in meters) from R as indicated by
fluorescent powder; DEST is the burrow or grid station at
which an animal was found the following morning. The actual
distance from R to DEST is shown in parentheses.
ID
DATE
R
ORI
DIST
DEST
F23
25
May
85
35
222
5.5
-
F30
25
May
85
13
125
11.9
-
it
14
Jul
85
81
302
9.9
71 (36.0)
F39
25
May
85
58
70
17.7
-
F47
15
Jun
87
6,2
47
20.0
6,0
Mil
25
May
85
10
162
15.2
10
M3 2
14
Jul
85
81
140
28.0
15 (45.0)
M3 9
14
Jul
85
22
108
19.2
23 (15.0)
M49
15
Jun
85
9,4
273
2.0
9,5 (10.0)
M54
15
Jun
85
9,1
250
15.0
-
M98
25
May
85
49
281
11.5
—

44
part of the night's activities. Because the fluorescent
powder yielded so little information regarding home range,
feeding behavior, or social activity, I concluded that it
was of little use with Podomvs at Ordway.
Trapping on Grids
Results of trapping on the three grids are shown on
Fig. 7, which includes data from all animals, including
transients caught only once or twice. A total of 18 animals
was trapped on the AC grids. I calculated persistence on
the grid (PQ) as the number of days from first to last
capture. Time spent by each individual on the grid is
represented by the horizontal bars; counting bars vertically
indicates the size of the known population per month. The
largest population occurred on the SL grid in the spring of
1987 and 1988, when there were 8 individuals or 8 ani¬
mals/ha. Excluding animals caught three times or less, mean
persistence time on this grid was 195.9 days for females and
137.2 days for males.
For 20 mice captured four or more times, I plotted the
capture locations and number of captures. To calculate the
home range area, I used the Exclusive Boundary Strip Method
(Stickel, 1954), in which I mapped the capture sites and
drew a boundary strip (equal to half the distance between
traps). I selected this approach because Stickel (1954)
compared the exclusive boundary strip, inclusive boundary

45
ID 1987 1988
M J J A S 0 M A M J J S N
ACI
TN 600
200 300 300 300 300
300
300 300 300 300 300
F89
(21)
F91
(75)
F92
(1)
F94
(1)
F95
(3)
F96
(21)
M94
(21)
M95
(21)
M96*
(21)
M99
(3)
F97
(3)
F85*
(1)
AC 11
TN
300 200 300 300
300 300 300 300 300
F88
M89*
M92
F87
M86
M84
(3)
(1)
(2)
(33)
(1)
(3)
Fig. 7. Trapping data for Podomvs on the ACI, ACII, and SL
grids. ID indicates sex and identification
number; asterisks indicate animals first captured
as juveniles; TN indicates trapnights per month;
and persistence on grid in days is shown in
parentheses.

ID
1987
1988
46
A M J
J A S 0 M A
M J J
A S N
SL
TN300 600 100
300 300 200 300 300
300 300 300
300 300
F10
(42)
F18*
(36)
F26*
(449)
F47*
(517)
F64
(364)
F61
(24)
Mil
(91)
Ml 6
(148)
M19_
(1)
M49
(91)
M54
(381)
M42
(289)
M98
(24)
M58
(1)
F42
(1)
F54
(74)
F58
(2)
M26
(1)
M53
(76)
M56
(44)
M59
(74)
M102
(156)
F35
(61)
M79
(1)
Fig. 7— continued

47
strip, and minimum area methods of calculating home range
area and she determined that the minimum area method under¬
estimated home range size by 64%, and the exclusive and
inclusive boundary methods overestimated home range size by
2%, and 17%, respectively.
On ACII only one individual (female 87 on Fig. 7) was
captured at least four times. However, her last two cap¬
tures occurred across the grid from the first three, and
this pattern suggests dispersal rather than a large home
range, so I did not include her in the analysis. Home range
data are summarized on Tables 5 and 6. Three animals were
marked on the grids as juveniles (gray pelage) and died or
dispersed off the grid at approximately 60 days of age. Two
more animals (female 26 and male 54 at SL) marked as juven¬
iles did not leave the grid. The male shifted his home
range, but I could not distinguish between juvenile and
adult home ranges for female 26, who subsequently appeared
to have an activity area larger than that of other females.
Seventeen adults trapped on the two grids (ACI and SL) had
an average home range of 804 m2. In general, males (mean =
910 m2) appeared to have larger home ranges than females
(mean = 683 m2), but the difference was not significant
(Mann-Whitney test at p = 0.01).
To determine whether ranges of females and males over¬
lapped, I plotted home ranges of Podomys caught during May
1987 on ACI and May 1987, September 1987, and May 1988 on SL

48
TABLE 5. Home range data for 20 Podomys trapped on two
grids. Id is the identification number, NG is the number of
captures, P0 is the persistence on the grid in days, A is
the area of the home range in m2 for juveniles (Aa) and
adults (AjJ .
Grid
Id
Sex
Ng
pG
AC I
89
F
6
21
700
AC I
91
F
7
75
500
AC I
96
F
4
21
400
AC I
94
F
6
21
916
AC I
95
M
5
21
750
AC I
96
M
5
21
800
SL
61
F
5
24
1000
SL
10
F
6
42
500
SL
18
F
6
36
600
SL
26
F
17
449
1350
SL
47
F
12
517
300
SL
54
F
6
74
300
SL
42
M
8
289
1850
SL
49
M
10
91
1200
SL
54
M
16
381
1093
200
SL
16
M
9
148
800
SL
53
M
6
76
600
SL
56
M
7
44
500
SL
59
M
8
74
1000
SL
102
M
10
156
400

49
TABLE 6. Capture data for 36 Podomvs at Anderson-Cue. ID
indicates sex and identification number; YR is year of first
capture; AGE is age at first capture (juvenile, subadult, or
adult); N is number of captures at grids (NG) , burrows (NB) ,
and total (NT) ; P is persistence, in days, on grids (PG),
burrows (PB), and total (PT) . Asterisks indicate trap
mortality.
ID
YR
AGE
Nb
PB
ng
Pc
Nt
PT
F83
88
J
1
1
1
1
F84
88
J
3
3
3
3*
F85
88
J
1
1
1
1
F86
88
A
7
109
7
109
F87
88
SA
2
2
5
33
7
33
F88
87
A
1
1
3
3
4
9
F89
87
A
6
21
6
21
F90
87
SA
1
1
1
1
F91
87
SA
7
75
7
75
F92
87
A
1
1
1
1
F93
87
A
1
1
1
1
F94
87
A
7
164
1
1
8
164
F95
87
A
1
1
3
3
4
21
F96
87
A
1
1
4
21
5
40
F97
87
A
9
171
3
3
12
193
F98
87
J
1
1
1
1
F99
87
A
1
1
1
1
F100
87
SA
4
124
4
124

50
TABLE 6 continued
ID
YR
AGE
Nb
PB
Ns
pG
N_
T
PT
M80
88
SA
3
216
3
216
M84
88
A
2
39
2
3
4
39
M85
88
SA
3
101
3
101
M86
88
SA
1
1
1
1
M87
88
SA
2
2
2
2
M40
87
A
1
1
1
1
M88
87
J
1
1
1
1
M89
87
J
1
1
1
1
M90
87
J
1
1
1
1
M92
87
A
3
48
2
2
5
48
M93
87
SA
3
5
3
5*
M94
87
A
2
106
6
21
8
106
M95
87
A
5
21
5
21
M96
87
J
5
21
5
21
M97
87
A
4
150
4
150
M98
87
?
1
1
1
1
M99
87
SA
1
1
3
3
4
21
M100
87
A
1
1
1
1

51
(Fig. 8), the three months during which populations appeared
to be the largest (Fig. 7). Generally, ranges of males and
females overlapped, but ranges of adult females did not
(except occasionally with juveniles or young adults), a
pattern which suggests intra-sexual territoriality (Kau-
fmann, 1983). A female juvenile (female 18) overlapped with
the ranges of an adult of each sex (Fig. 8A). Female 26,
the one with the unusually large range, overlapped somewhat
with female 61 in September 1987. The only males that
overlapped were three males at SL (Fig. 8C), two of which
possibly were siblings.
Trapping at Burrows
With the data from trapping at burrows, I estimated
home ranges by mapping movements of adults trapped more than
four times in a year on AC and SL, using the exclusive
boundary method (Stickel, 1954) as I did for grids. For
burrows on the perimeter of the study site, I plotted a
boundary of 10 m (which I considered a conservative estimate
of distance moved beyond the burrow). All capture points
were weighted egually, but in a few instances I omitted
points that suggested unusual forays outside the usual home
range of the animal. I then measured the home range area
with a Lasico Compensating Planimeter.
Seventy-five home ranges for 62 animals who fit the
above criteria, including three females on AC, are shown in

52
Fig. 8. Home ranges of Podomys trapped on the ACI grid in
May 1987 (A) (n = 6 mice, 600 trapnights) and on
the SL grid in May 1987 (b) (n = 8, 600 trap-
nights), May 1988 (c) (n = 9, 300 trapnights), and
September 1987 (D) (n = 7, 300 trapnights). Dots
represent the outer boundary of grid. Solid lines
represent home ranges of females, dotted lines
those of males.

53
Fig. 9. Home ranges for adult males frequently overlapped,
but those of females did not; two SL females who appear to
overlap in 1988 actually were separated temporally. This
separation of home ranges, similar to that observed on
grids, suggests intra-sexual territoriality. Animals who
survived more than one year (e.g., SL male 11 in 1985-87)
showed fidelity to the same general area every year. Home
ranges varied in size from 210 m2 (for SL female 26, cap¬
tured at a single burrow 6 times in 1987) to 11007 m2 (for
SL male 11 in 1986). The average home range size for 35
females was 2601 m2 (SD = 1315.1) and for 40 males was 4042
m2 (SD = 2269.4); home range size of the two sexes differed
significantly (Mann-Whitney U = 2.95, p = 0.01). For the SL
data I also compared areas in 1983-85 with those of 1986-88
to see whether home range size changed after the prescribed
burn in 1985, but I found no significant difference.
Capture data for the 62 mice whose home ranges were
mapped are shown on Table 7. Females 86, 94, and 97 (1987)
were from AC; the remaining animals were on SL. Each animal
was trapped from 5 to 22 times during the year in question,
and almost all mice utilized more than one burrow (mean
number of burrows = 3.8). Estimates of persistence (PB)
were calculated by counting the number of days between first
and last captures spent by the animal in the area shown on
the figure. These estimates of persistence are not com¬
parable across years because trapping effort varied each

Fig. 9. Annual home ranges of 62 Podomys on the SL
sandhill, 1983-1988. Dots represent tortoise
burrows where mice were trapped. Solid lines
indicate home ranges of females, dashed line for
males; a) 1983, 52 burrows; b) 1984, 61 burrows;
c) 1985, 89 burrows; d) 1986, 103 burrows;
e) 1987, 131 burrows; f) 1988, 136 burrows. Bar
represents 20 m.

55
Fig.
9a

56
85
Fig. 9b

57
100
Fig. 9c

58
Fig. 9d

59
Fig. 9e.

60
Fig.
9f

61
TABLE 7. Capture data for the 62 adults whose home ranges
are shown on Fig. 9. ID is the sex and identification
number of each mouse; YEAR is the year for which the home
range was calculated; N is the number of captures, PB is
persistence (in days); #B is the number of burrows where
each animal was caught; and asterisks denote animals present
when trapping ceased in November 1988.
ID
YEAR
Nb
PB
F5
1983
5
93
F8
6
71
F14
6
71
F27
5
114
F49
5
117
M2 9
5
79
M3 3
5
79
M47
5
99
M48
5
100
M53
5
79
F23
1984
8
125
F29
11
139
F34
5
56
F38
5
49
F43
15
222
M25
6
135
M27
5
61
M28
5
63
M3 9
14
171
#
B
2
3
2
2
4
2
2
2
1
2
2
5
2
2
3
3
3
2
5

62
TABLE 7--continued
ID
YEAR
Nb
PB
M4 8
12
211
M85
6
99
F23
1985
14
212
F39
12
84
F30
8
183
F97
8
185
F100
22
265
Mil
16
253
M3 2
13
183
M3 9
18
265
M98
13
265
M100
7
100
F89
1986
5
49
F93
5
36
F94
8
103
F95
6
107
Mil
19
253
M72
5
68
M73
7
79
M76
11
133
M80
12
103
M88
16
259
M97
5
101
8
3
3
4
5
3
5
5
6
7
6
5
2
3
4
2
8
3
3
7
7
4
3

63
TABLE 7—continued
ID
YEAR
nb
PB
M9 8
5
171
3
F26
1987
6
78
1
F47
8
113
2
F61
7
243
3
F68
6
170
5
F72
5
125
3
F86
7
109
4
F94
7
164
6
F97
9
171
8
Mil
8
143
5
M16
7
233
3
M17
5
193
2
M42
5
31
5
M57
9
242
3
M65
7
213
4
M80
9
201
5
M98
5
218
4
F10
1988
11
227*
4
F26
14
193
6
F36
5
94
3
F37
9
121
5
F46
10
151
5
F47
11
193
2

64
TABLE 7—continued
ID
YEAR
Nb
PB
#B
F49
8
148*
3
F52
11
151
5
M20
8
151
4
M2 2
8
151
7
M2 5
7
132
4
M3 4
6
104
4
M3 7
9
148*
5
M42
11
198*
7
M51
10
151
4
M102
13
198*
2

65
year; nor do they represent longevity because captures of
juveniles and subadults (or captures as adults in years
other than those shown) are excluded. The PB values are
useful as minimal estimates of persistence on the sandhill.
Conclusions
Fluorescent powder was not a useful method of determin¬
ing the home range of Podomys on Ordway. Powder trails did
not indicate the complete area traveled by mice even during
one night. Although some data regarding substrate use or
social interactions were gathered, considerable time and
effort were required. The technique proved successful with
smaller Peromyscus (Lemen and Freeman, 1985, 1986), so I
suspect the failure at Ordway was not related to the body
size or biology of Podomys, but possibly to density or
structure of vegetation.
Trapping on grids provided estimates of home range size
and persistence, as well as demonstrating the exclusivity of
home ranges of adult females. There also was evidence that
males had larger home ranges that occasionally overlapped
with each other and with females. However, estimates of
home range areas obtained from trapping at burrows suggest
that estimates from grids were too small. Evidently the
"standard" mammal-trapping grid ( 10000 m2 with 10-m trap
intervals) was insufficient to measure home ranges of Podo-
mys in sandhills. Doubling the trap interval to 20 m would

66
produce a 10 row by 10 column grid covering 40000 m2, which
would encompass mean home ranges estimated at burrows for
females (2601 m2) and males (4042 m2) , might be more useful.
No fewer than 100 trap stations should be used. However,
one should keep in mind that increasing grid size on some
sandhills will spread the grid into neighboring habitats,
and that trap success on the grid will be smaller than on
the burrows.
Trapping at burrows produced larger estimates of home
range size than I expected, but the means are not unreason¬
ably large when compared with other values reported for
Peromyscus♦ Stickel (1968) reported that home ranges of
Peromyscus varied from 0.1 to 10 acres. Hoffmeister (1981)
reported mean home ranges for female (8290 m2 +/- 2688) and
male (10465 m2 +/- 4043) P. truei in pinon-juniper habitat,
also using the exclusive boundary strip method. Although
the range of home areas of Podomvs varies considerably,
further research might show that my means are under¬
estimates, based as they are on many individuals located on
the edges of my study sites. I also have no measure of
vertical habitat use; for example; it would be interesting
to determine whether the mice occupy a larger home range
volume in years when acorns are produced.
In analyses of both burrow and grid data, I omitted
animals that were recaptured less than three times. A pos¬
sible consequence of this approach is that several animals

67
might occupy the same space. However, the fact that both
sets of data show abutment of home ranges of adult females
seems good evidence of mutually exclusive home ranges, which
suggests that females might be territorial.

DIET
Field Observations
Almost nothing is known about the natural diet of
Podomys. Merriam (1890) reported an observation that mice
in the southeast ate the seeds of "scrub-palmettoes." Layne
(1970) and Humphrey et al. (1985) suggested that acorns
might be a major food source during mating years, based on
the association of P. floridanus with habitats with Q.
laevis and other oaks. Milstrey (1987) reported that captive
Podomys ate engorged soft ticks (Ornithodoros turicata
americanus) that parasitize gopher frogs and tortoises,
raising the possibility that these mice prey on other resi¬
dents of Gopherus burrows. Other foods presumably include
insects, seeds, nuts, fungi, and other plant material
(Layne, 1978) .
Five field observations at SL provided additional
dietary information. An adult male caught and released on
the grid (5 May 1988) readily ate a cricket (Gryllinae) I
offered to him. On 15 June 1987 at approximately 0245 EST,
an adult female (who had been trapped, powdered, and re¬
leased) caught a small moth and consumed all but the wings.
On 9 May, an adult female just released from a trap ate a
68

69
young shoot of Smilax auriculatus. On 20 July 1987, at
burrow #15, I also observed an adult male take a fruit of
the flag pawpaw (Asimina incarna) (Jones, 1989). After
being released from the trap, this mouse climbed 30 cm up a
plant on the apron of the burrow, and knocked a ripe pawpaw
to the ground. He then ran down, picked up the pawpaw, and
carried it 11.4 m to burrow #71. In July 1988 another mouse
took a pawpaw, again at SL. A female released from a trap
at a burrow hid in a hole at the base of a turkey oak ap¬
proximately 5 m NW of the burrow. In a few minutes she left
the hole, picked up a small pawpaw fruit (A. incarna), and
carried it back to the hole and nibbled on it.
Predation and dispersal of Asimina fruits have not been
well studied. Willson and Schemske (1980) reported opossums
and raccoons sampling unripe fruits of A. triloba in Il¬
linois. Norman and Clayton (1986) suggested that small
mammals probably help disperse Asimina in Florida and re¬
ported an observation of Gopherus eating ripe fruits. Human
consumption of A. incarna was reported by Bartram (1791).
Whether Podomys disperse these fruits, and whether pawpaws
are a significant source of food, remains unknown. Willson
and Schemske (1980) measured a caloric content of 4.5 kcal/g
for fruits of A. triloba in Illinois; fruits ranged in size
from 2.1 to 48 g dry weight. Fruit set for A. pboyata, A.
pyqmaea, and A. triloba is less than 10% (Willson and
Schemske, 1980; Norman and Clayton 1986). To the best of my

70
knowledge, nutritional values and fruit set have not been
reported for A. incarna, the most abundant pawpaw on Ordway.
However, pawpaws certainly are the largest fruits produced
in the high pine; surely they represent a significant addi¬
tion to the siammer diet of Podomvs.
Laboratory Observations
The maintenance diet of captive animals consisted of
rodent chow (Wayne Rodent Blox) and water provided ad libi-
tem. This diet was supplemented with lettuce, carrots,
apples, strawberries, sunflower seeds, mixed bird seed,
oatmeal, rice, mealworms, crickets, and acorns (Q.
michauxi).
In an attempt to determine what mice might eat under
natural conditions, I occasionally offered animals fruits
and seeds collected at Ordway. All of these offerings were
eaten: acorns (Q. chapmanii. Q. qeminata. Q. hemisphaerica,
Q. laeyis, Q. mvrtifolia. and Q. nigral, pine seeds (P.
elliottii and P. palustris), blueberries (Vaccinium mvr-
sinites), deerberries (V. stamineum), gallberries (Ilex
glabra ^, blackberries (Rubus argutus), gopher apples (Lica-
nia michauxii), pawpaw fruits (A. incarna and A. pygmaea),
flowers of Stillingia sylvatica. and seed pods of Crotalaria
rotundifolia and Galactia elliotti. Seeds and stems of
unidentified grass species were shredded and incorporated

71
into the cotton nesting material, and probably eaten as
well.
Predation on vertebrates appears limited to stressful
conditions. Eisenberg (unpubl. data) trapped an adult male
on SL in 1983 who detached the posterior part of a juvenile
red rat snake (Elaphe guttata) that was caught half way in
the trap. The mouse ate about 3.5 cm of the snake to the
bone. An adult male caught on AC in October 1987 ate the
viscera of a juvenile also caught in the trap. I also lost
a litter of three young in captivity on the only occasion
that I had a male and two pregnant females in the same
aquarium.
In general, feeding behavior follows that described for
Peromyscus by Eisenberg (1968), in which the animal crouches
and manipulates the food with its forepaws. Larger items
such as pawpaws are held against the substrate. Seed pods
were opened by grasping the pod vertically, resting one end
on the substrate, chewing off one end of the pod, and open¬
ing the pod longitudinally along a suture. Larger foods,
such as turkey oak acorns, are dragged with the incisors;
smaller items are carried in the mouth. Some individuals
consistently took favored foods (such as acorns and sunflow¬
er seeds) to the corners of the aquarium and covered them
with kitty litter; I considered this caching behavior. Food
items and remains also commonly are found underneath nests.

72
This ready acceptance of a wide variety of foods im¬
plies that P. floridanus is an opportunistic feeder, like
Peromvscus. As King (1968) pointed out, we lack systematic
studies of taste discrimination and food preferences. For
Podomvs we also need basic studies (stomach and fecal ana¬
lyses) to learn more about the natural diet.
Acorn Preference Tests
I performed preference tests to determine whether
acorns of turkey oaks (Q. laevis) are preferred over acorns
of other species that are less abundant or produce fewer
acorns in the high pine. Such a preference might be a
partial explanation for the distribution of Podomvs in
sandhills at Ordway.
For feeding tests, an individual was placed in an
aquarium with clean kitty litter, nesting material, water,
and rat chow at least 24 hours before beginning the test.
To start a trial the chow was removed and three bowls (each
containing five acorns of a single species) were added at
about 1900 hours. Each acorn was marked and weighed and I
measured its height and width with dial calipers. No effort
was made to ascertain that all acorns in a bowl were identi¬
cal in size. Acorns that had weevil holes were not used,
and in a single trial all acorns either had caps or lacked
them. Approximately 12 hours later I removed the bowls,
acorns, and acorn fragments. I recorded whether acorns were

73
removed from bowls and whether they were opened and eaten,
opened and evidently not eaten, gnawed, or apparently un¬
touched. I controlled for location effects by shifting the
relative positions of the three types of acorns in each
trial.
Two unexpected difficulties arose. The first was the
unpredictability of the acorn supply, which made it impos¬
sible for me to run tests with identical species in 1988 and
1989. Additionally, the majority of acorns on the ground
already contained weevil larvae or were otherwise spoiled.
Secondly, I discovered that some of the acorns I tested were
spoiled (by insects or by mold) even though they lacked
holes. Possibly this spoilage is the reason that some hulls
were opened but the meat not removed. For this reason I
simplified analysis of results by examining whether acorns
were unopened or opened, regardless of whether any of the
meat appeared to have been removed. These data were then
ranked (rank 1 to the smallest number opened) and the Fried¬
man test (Conover, 1980) was used to test the hypothesis
that species of acorns were opened in equal numbers.
In 1988, I concentrated on determining which acorns
were eaten by Podomvs. I presented acorns from six species
of oaks (Chapman's (Q. chapmanii), live (Q. geminata),
turkey (Q. laevis), laurel (Q. hemisphaerica). myrtle (Q.
myrtifolia). and water oaks (Q. nigra)) to four captive
animals. Five of these species belong to the red-oak group,

74
which generally contains three or four times more tannin
(Briggs and Smith, 1989) than species of the white-oak group
(which includes Q. chapmanii). Each mouse was presented
with turkey oak acorns in combination with acorns from two
other species; two to four trials were run per animal for a
total of 12 trials. Although the sample is inadequate for
statistical analysis, I noted that the mice opened acorns of
all species, and in all but one trial turkey oak acorns were
opened in the smallest numbers. I then planned to test
animals with a combination of acorns from Chapman's, live,
and turkey oaks, the most prevalent Ouercus species on the
SL sandhill.
In 1989 Chapman's and live oak acorns were not avail¬
able, so I gathered acorns from Q. laevis and two different
trees of Q. hemisphaerica. one from the old pasture near
Ross Lake Bridge and the second from a hammock past Ander-
son-Cue. I expected that Q. laevis, the predominant acorn-
producing species on SL, would be preferred. I tested nine
animals, two trials each. For analysis I used results of
the first trial only; there was no difference in ranks of
first and second trials for any of these mice. Results
indicated that acorns were not opened in equal numbers
(Friedman test, T = 26.75, significant at p = 0.01). When
the Friedman test indicates rejection of the null hypothe¬
sis, multiple comparisons may be made to determine which
treatment is significantly different from the others

75
(Conover, 1980). For multiple comparisons at a significance
level of p = 0.01, acorns from Ross Lake (Q. hemisphaerica)
were opened significantly more often; differences between
the acorns from the hammock and from Q. laevis were not
significant. These results indicate not only that the mice
preferred laurel oak acorns over turkey oak acorns, but that
mice distinguished acorns from two different individuals of
Q. hemisphaerica.
The apparent preference for Q. hemisphaerica was sur¬
prising in that its acorns contain more tannic acid than Q.
laevis (Harris and Skoog, 1980). The laurel acorns were
smaller than the turkey oaks', so possibly the preference is
due to differences in handling time. Another possibility is
selection according to chemical composition other than
tannin content. Halls (1977) reported that Q. laurifolia
(= Q. hemisphaerica) had higher fat and carbohydrate content
than Q. chapmanii and Q. incana; unfortunately, values for
Q. laevis were not listed. Possibly my results were some
artifact of using captive animals. However, in the only
published study of acorn preference in Peromvscus, Briggs
and Smith (1989) found that five P. leucopus captured in
habitats lacking oaks consumed equal amounts of acorns from
species of the red and white oak groups. Peromvscus caught
in areas containing oak trees selected acorns of Ouercus
species found in their habitat, independent of fat, protein,

76
and tannin content. These results lead me to believe that
there is some real difference between the acorns I used.
Except for smaller acorns that occasionally were split
in half, acorns were opened consistently at the hilura (basal
scar). Although acorns typically were carried by the point
at the distal end, the mice never chewed open the hull
there. For small, round acorns a neat incision was made
around the scar; on more elongate nuts (Q. qeminata and Q.
laevis) the hull might be nibbled farther down the sides.
Caps, if present, were removed; there was no difference in
the numbers of capped and capless Q. laevis acorns opened.
Occasionally I found acorns opened in this manner on aprons
or near burrow entrances. Hulls of Q. laevis were found in
excavated burrows (Jones and Franz, in prep.) and E. Mil-
strey vacuumed remains of what appear to be Q. qeminata and
Q. laevis out of burrows on Roberts Ranch, Putnam County;
all of these hulls were opened in a manner consistent with
what I observed with captive animals. Small piles of emptied
hulls occasionally are found at the base of turkey oaks on
Ordway; however, it is possible that Glaucomvs or birds
might make these piles. Of the 450 acorns presented in 30
trials to 13 mice, 66% were removed from bowls, whether
opened or not.
I avoided use of acorns that appeared spoiled because
mice did not appear to eat blackened nutmeats. I did not
test preferences of sound acorns vs. acorns with weevil

77
larvae. On one occasion an adult female immediately ate a
larva from an acorn opened but not eaten by another in¬
dividual. Semel and Andersen (1988) suggested that such
differences in behavior might be due to mice being unable to
detect larvae in unopened acorns, or that individuals can
detect infestation but some individuals avoid infested nuts.
They thought that tooth marks on the hull and movement of
acorns might represent assessment.
Based on these results, I suggest that acorns of Q.
laevis are eaten but other species are preferred if avail¬
able. Turkey oaks are an unreliable food supply. Umber
(1975) noted acorn production of Q. laevis was low and
variable on his study site in Citrus/Hernando counties, and
Kantola and Humphrey (in prep.) found that trees on low
slopes at Ordway produced significantly more acorns than
trees on higher slopes. Layne (in press) correlated the
relatively greater abundance of Podomys in scrub and scrubby
flatwoods with higher and more consistent acorn production
than in the high pine. Proximity to Q. aeminata and other
acorn-producers might be a partial explanation for the
relatively higher and more constant population of Podomys at
SL than at AC and BP. If oak species in hammocks are more
reliable producers, the restriction of P. aossvpinus and
Ochrotomvs nuttali to hammocks at Ordway might be partly due
to the food supply. This leads to the hypothesis that
competition, possibly linked with other factors such as

78
predation, forces Podomys into the high pine when the other
rodents are present.
Supplemental Feeding Experiment
Are Podomys limited by food supplies? If food is a
limiting factor, mice should respond to extra food with an
increase in numbers, increased individual mass, and an
extended breeding season. A food limitation could partly
explain population sizes and distributions at Ordway. I
attempted an experiment of food supplementation using foods
favored by captive Podomys. The only supplementation ex¬
periment that has been done with the Florida mouse was that
by Young (described in Young, 1983 and Young and Stout,
1986) on two grids in sand pine scrub in Orange County.
Other rodents responded to the additional food, but Podomys
rarely appeared on Young's grids and failed to establish a
permanent population during the experiment, although the
species had been abundant on both grids before the study
(Young, 1983). She concluded that Florida mouse populations
were limited by factors other than food.
My experimental design consisted of three grids, two on
the AC and one on the SL sandhill. Trapping procedure was
described under Home Range. The last grid (SLII) was set on
the SL sandhill west of Breezeway Pond. Prior trapping at
burrows in each of these areas indicated that mice were
present. In April, May, and June of 1988 I trapped SLI for

79
800 trapnights to determine how many nights of trapping were
needed to obtain all animals on the site. Consequently,
SLI, ACI, and ACII were trapped for three consecutive nights
per month for a total of 300 trapnights monthly. On ACI a
mixture of sunflower seeds, mixed bird seed, and oatmeal was
provided for one year in seven chick feeders fitted with
glass jars. To eliminate non-target species, plastic buck¬
ets with two one-inch holes cut near the rims were upended
over the feeders and weighted down with a cinder block. I
checked feeders periodically and refilled as needed.
The results of this experiment are shown in Fig. 10.
Capture rates declined sharply on AC and were highly vari¬
able on SL. I had intended to provide food on a second grid
(SLII, west of Breezeway Pond). However, 208 trapnights on
this grid and surrounding burrows in May, July, and August
resulted in the capture of one Podomvs. The lack of success
on ACI led me to abandon SLII although I continued to pro¬
vide food on ACI until the end of the 1988 trapping year.
On the AC sandhill I continued to trap mice in small numbers
at burrows west and east of ACI. Clearly, mice were not
obtaining the food or some other factor kept them out of the
grid area in spite of obtaining food.
Maximum trapping success for a single night was 9%, but
success was usually much lower. Additionally, I captured
non-target species, including Sceloporus undulatus, Coluber
constrictor, and Glaucomvs volans. In spite of the low trap

Fig. 10. Minimum numbers of individuals (MNI) on the ACI, ACII, and SL grids
Arrows indicate beginning and ending of food supplementation on ACI

1987 1988
MONTH
GRID
GRID
GRID
oo

82
success I attempted to estimate the persistence (i.e., the
time between first and last captures) of mice on the three
grids (Table 8). Because these were not closed grids,
disappearance of an animal might be due to death, dispersal,
or to not entering traps.

83
TABLE 8. Persistence data for animals on grids. Data
presented are minimum number known alive (MNA), mean +/- SD
days
total
spent on a
numbers of
grid (maximum values
trapnights (TN).
in parentheses),
and
Grid
MNA
PERSIST
TN
AC I
12
16.00
+/-
20.86
(75)
4200
AC 11
6
7.17
+/-
12.68
(33)
3000
SLI
24
122.75
+/-
155.7
(517)
4100

ANALYSES OF VEGETATION
Sampling Technique
Can the local distribution of Podomys be predicted on
the basis of vegetation structure or species composition?
Vegetation was sampled in circular quadrats (radius = 10 in,
area = 314.159 m2) set with tortoise burrows as center
points. In the absence of large numbers of trees, circular
quadrats are laid out easily using a center pole at the
burrow and a rotating radius line; forestry flags were used
to delineate the quadrat. Quadrat size was based on prelim¬
inary estimates of activity areas obtained by dusting Podo-
mvs with fluorescent powder. Although one animal ranged at
least 28 m from his burrow in one night (see Home Range
above), I decided a radius of 10 m was sufficient to contain
significant numbers of plants and small enough to allow me
to feasibly obtain counts and identifications of plant
species. When I began these samples in 1985 I was uncertain
that burrows were distributed randomly, so point-quarter
sampling seemed inappropriate. Without knowing the shape of
Podomys activity areas, I felt that line-intercept sampling
and belt transects would be less representative of areas
encountered by mice than circular quadrats.
84

85
Sandhills on the Ordway Preserve have been subjected to
logging, grazing, suppression of fire, and prescribed burn¬
ing on different rotations. Cessation of fire, or the
frequency of prescribed burns and other activities, affect
plant species composition and density on sandhills (e.g.,
Myers, 1985; Gates and Tanner, 1988). To determine how
burrow use changes with successional changes in vegetation,
I examined quadrats at SL in 1985, 1987, and 1988. SL has
been burned and the pines bear turpentine scars. Trapping
effort was equal among these burrows within each year but
not among years. Initially 29 quadrats were examined; this
number was reduced to 27 after a large oak fell over one
burrow and a clearcut was performed at a burrow under a
powerline. To see how Podomvs behaved at burrows on a
sandhill where logging occurred, I set quadrats at 15 bur¬
rows at BP in 1988.
I selected burrows so as to include burrows of all
three classes of tortoise activity (active, inactive, or
old, as defined by Auffenberg and Franz, 1982). Within each
quadrat, I counted stems of all woody species (Table 9) in
three size categories (tree = dbh > 7.6 cm, sapling = height
> 1 m and dbh < 7.6 cm, and seedlings = height < 1 m). I
noted the presence or absence of woody debris, including
fallen branches or trees, or rotting logs. I estimated the
per cent ground cover of bare ground, wiregrass, and gopher
apple using five categories (I = 0-10%, II = 10-25%,

86
TABLE 9. Habitat variables measured in each quadrat. Prime
signs indicate variables used in discriminant analyses.
Variable (abbreviation)
Description
Location (LOC)'
BP or SL sandhill
Year (YR)'
1985, 1987, or 1988
Burrow (BUR)'
Identification number
Orientation (ORI)'
Compass reading away from
entrance
Sand pine (SP)'
Total stem count (P. clausa}
Slash pine (HP)
" (P. elliotti}
Longleaf pine (LP)'
" (P. palustris}
Loblolly pine (YP)
" (P. taeda}
Bluejack oaks (BO)'
" (0. incana}
Turkey oaks (TO) '
" (0. laevis}
Laurel oaks (LA)
" (0. laurifolia}
Post oaks (PO)
" (0. stellata}
Live oaks (LO)'
" (0. aeminata}
Persimmon (PER)'
" ÍD. virainiana}
Pawpaws (PAW)'
" (Asimina spp.}
Rosemary (RO)'
" (C. ericoides}
Sumac (SUM)'
" (R. copallina}
Blueberry (VAC)'
" íV. mvrsinites}
Debris (DEB)'
Presence or absence of fallen
logs or branches in each
quarter of quadrat (4 indi¬
cates presence in all 4 quar¬
ters )

87
TABLE 9—continued
Variable (abbreviation)
Description
Bare ground (GRO)'
Average of categorical es¬
timates of % of bare ground
coverage in each quarter of
quadrat
Bare ground (2GR0)'
Summation of categorical
estimates of bare ground
coverage per quadrat
Wiregrass (WIR)'
Average of categorical es¬
timates of % of coverage by
wiregrass of each quarter of
quadrat
Wiregrass (2WIR)'
Summation of categorical
estimates of wiregrass cover¬
age per quadrat
Gopher apple (GA)'
Average of categorical es¬
timates of % coverage by
Licania of each cruarter of
quadrat
Gopher apple (EGA)'
Summation of categorical esti¬
mates of Licania cover per
quadrat
Legumes (LEG)'
Total number of species of
Fabaceae
Herbs (HERB)'
Total number of all non-woody
species

88
III = 25-50%, IV = 50-75%, V = 75-100%). I grouped species
of grasses into five categories (I = Andropoqon spp.;
II = Panicum hemitqnum, III = Panicum (DicantheliunU spp.,
IV = Paspalum setaceum. V = Gvmnopoqon spp.) and sedges into
two ("small", such as Bulbostylis warei, and "large", such
as Cyperus retrorsus), but did not identify each species. I
identified and counted the number of herbaceous species
(excluding grasses and sedges), but I did not count stems.
All plant species (Appendix) were identified using the plant
list compiled for the preserve by David W. Hall and Richard
Franz.
I plotted a species-area curve (cumulative number of
species vs. cumulative number of samples) for the 1985 data
from SL; the curve levelled off after the fifteenth sample,
and I concluded that the number and size of quadrats were
adequate. I believe smaller sample areas would be inap¬
propriate for two reasons: Hansen (1964) noted that only 8
of 24 burrows he examined had turkey oaks and pines within a
radius of approximately 305 cm around the burrow entrance,
so larger areas are required to represent trees in the
vicinity. Secondly, some quadrats I examined were located
at burrows with large aprons, which are disturbed areas that
might be favored by certain animals such as Tantilla relic¬
ta , Eumeces eqreqius. and Neoseps revnoldsi (Campbell and
Christman, 1982); it is unknown whether these species affect
plant distribution on the apron. I suspect that burrow

89
construction also has some impact on density or heterogene¬
ity of local plant species (perhaps similar to that des¬
cribed at badger excavations by Platt in 1975), although
such an effect has not been documented in the literature for
Gopherus. Certainly the abundance of some high pine species
(such as Eupatorium compositifolium, according to Grelen,
1962 and Hebb, 1971) is affected by other disturbances such
as chopping or burning.
Results and Discussion
How did vegetation on the SL plots change between 1985
and 1988? On the burned plots, I expected increased stem
counts of woody species and increased numbers of herbaceous
species after the 1985 burn. On the unburned plots, I
expected little or no change in species composition or
density over the three-year period. Frequencies and den¬
sities of woody species are shown in Table 10. Two changes
were quite evident. Following the fires, particularly the
first burn in 1985, numbers of stems of Ouercus and Asimina
increased. Secondly, mortality of longleaf pine seedlings
(P. palustris) obviously was quite high in some areas on
both burned and unburned sites due to pine rust in 1986.
Stem counts of P. palustris were much lower in 1987 and stem
counts of other woody species, such as sumac, increased in
the affected areas.

90
TABLE 10. Frequency (% = percent of quadrats in which a
species occurred) and density (# = average number of stems
per 314 m2 quadrat) of woody species identified at SL.
YEAR
Burned plots (n =
22)
1985
% #
1987
% #
1988
% #
P. clausa
31.8
0.6
18.2
0.2
27.3
0.6
P. oalustris
100.0
151.6
100.0
66.9
100.0
81.4
0. aeminata
95.5
66.5
95.5
157.5
95.5
173.8
0. incana
100.0
88.4
100.0
163.2
100.0
159.9
0. laevis
100.0
97.8
100.0
174.6
100.0
156.3
0. laurifolia
13.6
1.5
13.6
3.1
13.6
8.2
Asimina sdd.
100.0
40.6
100.0
44.6
100.0
48.7
Ceratiola
31.8
0.9
18.2
0.6
27.3
0.6
Rhus
90.9
86.9
100.0
104.2
95.5
82.9
DiosDvros
95.5
34.2
100.0
37.1
100.0
44.8
YEAR
Unburned plots (n =
5)
1985
1987
1988
%
#
%
#
%
#
P. clausa
0
0
0
0
0
0
P. oalustris
100.0
189.4
100.0
141.8
100.0
117.6
0. aeminata
100.0
20.0
100.0
28.4
100.0
28.4
0. incana
100.0
33.8
100.0
36.4
100.0
43.6
0. laevis
100.0
153.0
100.0
204.6
100.0
219.8
0. laurifolia
0
0
0
0
0
0
Asimina spp.
100.0
14.0
100.0
25.4
100.0
25.4
Ceratiola
20.0
0.8
20.0
0.8
20.0
0.8
Rhus
40.0
20.4
40.0
41.2
40.0
28.0
Diosovros
100.0
10.0
100.0
27.4
100.0
31.2

91
How did vegetation on the BP and SL quadrats differ?
SL has more species considered "characteristic" of the high
pine environment; the less characteristic species, such as
Ouercus laurifolia, Q. mvrtifolia, Prunus serótina, and
Ervthrina herbaceae. were found on quadrats at the periphery
of the sandhill. The perception that there are fewer long-
leafs at BP is valid (Table 11). Also, P. clausa and Cera-
tiola are absent from the BP quadrats, reflecting the low
numbers of these species on the BP sandhill. Specimens of
P. elliotti and P. taeda, both absent on SL, grow at burrows
located on the periphery of the unburned BP site and old
pasture. Numbers of turkey oak stems are much greater than
on SL and Q. stellata is present. Densities of persimmon
(Diospyros virqiniana^ are somewhat lower on BP; however,
the unburned BP quadrats were the only ones containing
sapling-sized persimmons. Laessle (1942) and Grelen (1962)
noted that Diospyros become more abundant when fires are
suppressed and reach greater heights (> 2 m) following dis¬
turbance (e.g., chopping) than persimmons on undisturbed
sites. The saplings at BP probably reflect a history of
fire suppression following logging or some other major
disturbance. On BP I observed a few forbs (e.g., Chamaesvce
cordifolia. Convza canadensis. and Rudbeckia hirta) that are
associated with disturbed sites, according to Wunderlin
(1982); these species were absent or less abundant on SL.
However, when I compared the mean numbers of legume species

92
TABLE 11. Frequency (% = percent of quadrats in which a
species occurred) and density (# = average number of stems
per 314 m2 quadrat) of woody plant species identified at BP
in 1988.
Burned plots
(n = 9)
Unburned plots
(n = 6)
%
#
%
#
P. elliotti
0
0
16.7
0.3
P. palustris
11.1
0.2
50.0
2.0
P. taeda
0
0
16.7
0.2
0. aeminata
66.7
37.1
83.3
56.3
0. incana
66.7
8.7
83.3
20.0
0. laevis
100.0
521.7
100.0
257.2
0. laurifolia
0
0
16.7
0.5
0. stellata
55.6
18.3
33.3
9.3
Asimina sop.
100.0
47.4
100.0
19.0
Rhus
44.4
8.9
66.7
20.7
Diosovros
88.9
33.8
100.0
27.8

93
and herbaceous species on burned quadrats at BP and SL (1988
samples), I found no significant differences; nor did the
unburned quadrats differ significantly in these two parame¬
ters . Stem counts of woody species on burned and unburned
plots on BP were significantly different (X2 = 461.03,
highly significant at p = .005).
Can numbers of Podomys at particular burrows be pre¬
dicted based on local vegetation? A univariate analysis was
performed to test whether the minimum number of individual
mice (MNI) or total number of captures (ALL) could be pre¬
dicted using variables listed in Table 9. Then a second run
was performed using only the 21 parameters shown on Table
12. Because MNI and ALL generally were small, these vari¬
ables were set to 1 or 2 (if ALL <= 5, then ALL = 1, and if
ALL > 5 then ALL =2; if MNI = 0, then MNI = 1 and if MNI >
0, then MNI = 2). The null hypothesis was that there was no
difference between burrows used by mice (MNI > 0) and avail¬
able habitat, i.e., burrows not used by mice (MNI = 0).
The parameters I used were somewhat useful predictors
of ALL (Table 12), but did not predict MNI as accurately.
Only four variables (TO, LL, PER, and HERB) had significant
ability (p < 0.05) to predict ALL. Results of discriminant
functions analyses also indicated that these four variables
were the most significant. Captures were negatively corre¬
lated with stem counts of turkey oaks and longleaf pines,
and positively correlated with stem counts of persimmon and

94
Table 12. Habitat profile of Podomvs at all BP and SL
burrows where vegetation was sampled. Abbreviations are
given in Table 9. Univariate F values are given for anal¬
yses between habitat variables and ALL or MNI. N = sample
size. *p < 0.05, **p < 0.01, ns not significant.
VAR
ALL
MNI
N
89
89
LOC
0.46
0.02
YR
0.12
4.08*
BUR
0.19
0.15
ORI
0.74
0.15
TO
4.62*
0.41
BO
0.70
0.81
LO
3.45
0.13
LL
4.13*
0.90
PER
10.20**
0.02
SUM
0.47
1.73
PAW
0.83
1.10
DEB
0.18
0.87
ZGRO
0.48
0.53
GRO
0.71
2.36
2WIR
0.30
4.82*
WIR
0.06
4.31*
EGA
1.31
1.41
GA
0.12
5.57*
VAC
1.78
0.62
LEG
1.01
0.25
HERB
5.30*
0.83
Wilks' Lambda
0.58
0.70
X2
41.12
26.91
P
★ ★
ns

95
numbers of herbaceous species. Stem counts and herbaceous
diversity are affected by fire regime; the negative
correlation with oaks and pines is consistent with the
association of tortoise burrows with open canopy. Most of
the other oaks and all the persimmons (except some on un¬
burned quadrats at BP, where ALL = 0) were too small to pro¬
duce fruits. With the discriminant functions analysis, I
classified burrows to the two categories of total number of
captures (ALL) (Table 13). With these vegetation variables,
92% of the burrows in ALL <= 5 and 83% of the burrows in ALL
> 5 were classified correctly. Using MNI instead of ALL,
only 77% of the samples in MNI = 0 and 71% of the samples in
MNI > 0 were classified correctly. I repeated the dis¬
criminant function analysis using 9 parameters (TO, LO, LL,
PER, GRO, WIR, GA, LEG, HERB) representing stem counts of
woody species with F > 3.0, ground cover characteristics,
and numbers of herbaceous species. With this group of
variables, 70% of the burrows in ALL <= 5 and 66% of the
burrows in ALL > 5 were classified correctly. Therefore it
seems these variables are more useful for predicting rela¬
tive abundances of Podomys. than for predicting the presence
or absence of mice.
I thought it surprising that Podomys did not show a
closer correlation with indices reflecting ground cover. I
also ran a stepwise multiple regression, using the R2 cri¬
terion to determine the best-fitting regression equation for

96
TABLE 13
cording
of total
. Number (%) of vegetation samples
to discriminant function analysis to
number of captures (ALL) of Podomvs
reassigned ac-
two categories
TO:
ALL <= 5
ALL > 5
FROM:
ALL <= 5
65(92%)
6(8%)
ALL > 5
3(17%)
15(83%)

97
the two categories of ALL and the independent variables.
For the BP samples, the best equation (R2 = 0.97) was ob¬
tained using TO, BO, LO, PER, SUM, PAW, DEB, 2GR0, ZWIR,
EGA, LEG, and HERB. These results must be interpreted
cautiously (N = 15, df = 2). For the SL samples, the best
equation is obtained if LP also is included (R2 = 0.85,
0.49, and 0.34 for 1985, 1987, and 1988 samples respective¬
ly)-
Although BP and SL are both "high pine" or "turkey
oak-longleaf pine sandhills", the vegetation on the two
sites was not the same. If I had measured abundances of
wiregrass and other non-woody species the difference would
be more striking. Laessle noted for sandhills (1942:8),
"Among the most striking vegetational characters . . . are
the small but distinct floristic differences which they show
when compared to corresponding communities of the western
side of the ridge in central Florida." Of all the Ordway
sandhills on which I have found Florida mice (Anderson-Cue,
Blue Pond, Longleaf Pine, Rowan, and Smith Lake), no two
appear alike in spite of their similar classification. More
long-term studies that examine plant-animal interactions are
needed to compare the ecology of Podomvs on different sand¬
hills.
The burned part of BP appears to be developing into an
oak savanna dominated by Q. laevis; given the small number
of adult longleafs and the absence of a seed source, it is

98
unlikely that burning will create a longleaf pine savanna,
unless pines are planted (Platt et al., 1988; Rebertus et
al., 1989). At SL there appear to be plenty of longleaf
pines in all age classes, so possibly adherence to the
three-year fire regime will eventually eliminate turkey
oaks. However, it is unclear what effect this management
policy will have on Podomvs. Individuals seemed to persist
longer on burned sites, and there was a correlation with
stem counts of both oaks (mostly seedlings and saplings) and
pines. If oaks represent an important food source, Podomvs
might persist on a longleaf pine savanna only in low num¬
bers. Additionally, I would expect the decreased abundance
and diversity of herbaceous species after the suppression of
fire (Hilmon and Hughes, 1965; Gates and Tanner, 1988) to
have a negative effect on mouse populations, although we
still know little about the Podomvs diet.
Tortoises themselves apparently respond to a combina¬
tion of physical and herbaceous features, in that they
prefer abundant ground cover, high herbaceous diversity,
sparse shrub cover, and an open canopy, rather than specific
plant associations (e.g., Umber 1975; Auffenberg and Franz,
1982; Cox, Inkley, and Kautz, 1987; Breininger et al.,
1988). Effects of Gopherus on local vegetation are not well
known. Major components of the tortoise diet include
Poaceae, Asteraceae, Fabaceae, Pinaceae, and Fagaceae (Gar¬
ner and Landers 1981; Macdonald and Mushinsky 1988).

99
Possibly tortoises disperse some food species; Laessle
(1944) noted abundant gopher apple seeds around burrow
entrances. As mentioned previously, the physical impact of
the burrow excavation on the local vegetation has not been
studied, although Kalisz and Stone (1984) noted that excava¬
tions by burrowing animals help mix nutrients in the soil.

PRESCRIBED BURNS
Materials and Methods
The high pine sandhills of Florida are pyrogenic com¬
munities that were subjected to frequent, cool fires.
However, aside from the work by Mushinsky (1985) and Gates
and Tanner (1988), little is known about the effects of
these fires on members of the sandhill fauna besides the
gopher tortoise. Taylor (1981) summarized literature re¬
garding effects of prescribed fires on small mammals in the
southeast. Although the Florida mouse is listed as a
threatened species (Layne, 1978), in only one study (Arata,
1959) were the effects of fire on Podomvs examined. Using
traplines, Arata trapped for 150 trapnights before the burn
and for 1800 trapnights beginning within two weeks after the
burn. His trapping success for Podomvs was 3% and 9%,
before and after the fire respectively.
In my study of habitat preferences by Podomvs, I had
the opportunity to experimentally determine the effects of
fire on Podomvs by trapping three sandhills (BP, LP, and SL)
before and after prescribed burns. All three sites are on a
three-year rotation of prescribed burns; BP is burned in the
spring and LP and SL are burned in the late summer. Each of
100

101
these sandhills included a small area not to be burned. BP
was last burned in 1985. In 1988 we planned to burn BP in
May, so I trapped for two months before and after the sche¬
duled date of the burn, at 26 burrows to be burned and 13
burrows in the unburned area across the road. On 24 May
there was a lightning strike north of the pond and the
Ordway staff spread fire to the rest of the sandhill; the
site continued to burn through 25 May. From 27 May through
November 1988, I trapped all the BP burrows for a total of
2442 trapnights.
I briefly worked on LP in 1987. I trapped at 18 bur¬
rows on the area not to be burned for 375 trapnights between
26 July and 1 November. I trapped at 24 burrows on the side
to be burned for 492 trapnights (318 trapnights before the
fire on 19 September 1987, and 174 trapnights afterwards).
SL was last burned in September 1982; we burned this sand¬
hill again on 10 September 1985. In 1985 I monitored 57
burrows on the area to be burned and 22 burrows on the area
not to be burned. During the year of the burn, I trapped on
SL for 3092 trapnights, including 2308 on the burned and 784
at the unburned burrows. Because I found trapping success
higher at burrows than on transect lines, I continued to
trap at burrows. However, comparisons among BP, LP, and SL
must be made cautiously. Not only did the number of burrows
(hence, trapping effort) vary, but the sandhills differed in

102
area and seasonality of burns, and the unburned areas were
smaller than the burned.
To analyze trapping data, I followed Buech et al.
(1977) in calculation of a capture index (Cl), which equi¬
librates trapping effort to 1000 trapnights. They defined
Cl = number of individuals captured/1000 trap station
nights, where a trap station night consisted of two traps at
a station for 24 hours. For example, on the SL burn site:
1000 trapnights X 19 animals = 23.56 = Cl
805 trapnights
I also calculated minimum numbers of individuals (MNI)
per 100 trapnights per hectare in an attempt to equilibrate
trapping effort and area. The G-test was used to test the
null hypothesis that equal numbers of mice were captured on
burned and unburned sites.
Description of Procedure for Prescribed Burns
Prescribed burning is a tool used by land managers to
sustain a particular type of forest or savanna. Frequently
the ultimate goal is to maintain populations of special
interest: game and forestry species or threatened and en¬
dangered species. For example, in 1982 the Forest Service
burned approximately 81000 ha in the Southeast Region alone
to enhance habitats used by the red-cockaded woodpecker
(U.S. Dep. Agrie., 1984). Given these goals, it is not
unexpected that the majority of studies that examine how

103
animals are affected by fire have concentrated on one spe¬
cies of interest, such as quail or white-tailed deer, rather
than examining effects on less conspicuous species or on the
fauna as a whole.
Humphrey et al. (1985) summarized the management his¬
tory of the Ordway Preserve. Some sandhills on the proper¬
ty, intended for use as unburned "controls", have been left
unburned since at least 1980 when the University of Florida
acquired the preserve. The remaining sandhills were placed
on a three-year regime of prescribed burns at various sea¬
sons . The management goal is to return these sandhills to
longleaf pine savannas, as they are believed to have existed
(according to Bartram, 1791) before the initiation of logg¬
ing, farming, and ranching in the area.
On the Ordway, we conducted prescribed burns using drip
torches. Fire lines are dropped along roads and fire lanes,
and fires burn into the wind. We raked debris away from
snags and longleaf pines scarred by turpentine operations.
This method produces a relatively cool fire.
Results and Discussion
Apparent densities of Podomvs fluctuated monthly on all
three sandhills (Figs. 11-13). Numbers of mice were more
constant at burned sites than unburned sites. Apparent den¬
sities at SL frequently were larger at the unburned burrows;
possibly this was a result of the disparity in size of the

Fig. 11. Apparent densities (expressed as minimum numbers of individuals/ha) of
Podomvs on the burned and unburned areas of the Blue Pond sandhill.

.02
.01
v -
M A M J J A S
1988
• burned
O unburned
* fire
iriímiiH^
O N
105

Fig. 12. Apparent densities (expressed as minimum numbers of individuals/ha) of
Podomvs on the burned and unburned areas of the Longleaf Pine
sandhill.

MNI / ha
3.0
1.0
• burned
O unburned
* fire
1987
107

Fig. 13. Apparent densities (expressed as minimum numbers of individual/ha) of
Podomys on the burned and unburned areas of the Smith Lake Sandhill.

MN I / ha
o
VO

110
two areas. During my fieldwork I noticed that all animals
caught on the unburned sites (on all three sandhills) were
captured at burrows on the periphery of the unburned area;
none were captured in the middle of the unburned sites. For
three months preceding and following the burns, trapping
success on the three sandhills ranged from 2.2 to 8.4%; no
significant differences before and after the fires were
noted. On the unburned sites, trapping success ranged from
1 to 3.4%, the highest being on the BP ecotone.
Significantly more mice were caught in burned areas
both before and immediately following a fire (Tables 14 and
15). Significantly more individuals were captured on burned
sites (Table 16) than at unburned burrows (G = 10.597, sig¬
nificant at p < 0.005). The smaller difference in CIs be¬
tween burned and unburned BP burrows was due presumably to
the fact that all animals caught on the unburned area were
captured at burrows on the periphery of the site with old
pastures. There were numerous tortoise burrows, oaks, and
open areas on this periphery. No animals were caught in the
middle of the unburned area. The relatively smaller Cl
(11.84) for the burned burrows at BP was possibly due to
differences in vegetation and past land use.
Prescribed burns appeared to have little or no im¬
mediate effects (i.e., three months following the burn) on
Podomvs populations. Some populations decreased but two or
three months later populations equaled or exceeded pre-fire

Ill
TABLE 14. Burrows on burned sandhills. Data obtained from
trapping Podomvs at n burrows subjected to prescribed burns
on the BP, LP, and SL sandhills. # is total number of in¬
dividuals caught.
Pre-burn
Post-burn
BP(n = 26)
Trapnights
416
676
#
10
8a
Total captures
24
28
LP(n = 24)
Trapnights
318
174
#
5
5b
Total captures
7
5
SL (n = 57)
Trapnights
558
805
#
15
19c
Total captures
44
37
aincludes 4 individuals captured before the burn.
bincludes 2 individuals captured before the burn,
“"includes 10 individuals captured before the burn.

112
TABLE 15. Burrows on unburned sandhills. Data obtained
from trapping Podomvs at n unburned burrows on the BP, LP,
and SL sandhills. # represents number of individuals caught.
BP
LP
SL
n
13
18
22
Trapnights
596
375
497
#
6
4
6
Total captures
24
6
9

113
TABLE 16. Comparison of capture indices for Podomvs at
burrows on burned and unburned sandhills (after Buech et al.
(1977)). The burn/unburned column shows that the numbers of
individuals captured in burned areas average 194% of those
at unburned burrows.
burn Cl
unburned Cl
burn/unburned %
BP
11.84
10.08
117%
LP
28.75
10.68
269%
SL
23.56
12.06
195%

114
levels (Figs. 11 and 13, Table 17). Bock and Bock (1983)
suggested that fires in pine forest might increase the
quantity or quality of food supplies, thus producing a
temporary increase in the number of deer mice captured after
a fire. Whether this is what happens on the sandhills is
unknown. After the burns on my three sites, half of the
mice captured after the fire (all adults) had not been
caught before the fire; these animals might be dispersers
from surrounding, unburned areas. Layne (1974) and Martell
(1984) both suggested that burned areas might act as "dis¬
persal sinks". Tevis (1956b), Hatchell (1964), Ahlgren
(1966), Sims and Buckner (1973), and Krefting and Ahlgren
(1974) documented dispersal of Peromvscus onto burned areas.
Kaufman et al. (1988) suggested that odors from burned areas
might stimulate immigration of P. maniculatus from subop-
timal habitats.
To examine the effects of burns on home ranges, I
arbitrarily selected animals who were captured at least five
times, on the premise that two or three captures do not
reveal significant information regarding size of home ran¬
ges. For the 13 animals thus qualified, activity areas did
not significantly shift or change in size. At SL, the
animals caught more than five times appeared to be con¬
centrated on the periphery of the study area, near the xeric
oak hammocks. Home ranges for eight animals on SL are shown
on Fig. 14. The female (#70) captured in the middle of the

115
TABLE 17. Comparison of capture indices for Podomys at
burrows on 3 burned sandhills (after Buech et al. (1977)).
Pre-burn Cl
Post-burn Cl
Before/after
(%)
BP
24.04
11.83
203%
LP
15.72
28.74
54.7%
SL
26.88
23.60
114%

Fig. 14. Home ranges of eight Podomys on the SL sandhill before (stippled) and
after (hatched) the 1985 fire. Dots represent burrows where animals
were captured; asterisk indicates burrow used by both males 32 and 39.

117
O
o
z
?30

Fig. 14—continued

GO

119
burned site crossed the road to the unburned area between 15
and 22 September; on 13 October, the date of her last cap¬
ture, she had returned to a burrow on the burned side. This
was my only observation of an animal moving from burned to
unburned burrows after a fire, and she might have moved for
reasons unrelated to the burn. For those animals that
continued to be captured in 1986, no large shifts in ac¬
tivity area occurred. At BP, five mice were captured at
least five times in the two months before and after the
fire. Likewise, no major changes in activity areas were
observed. At LP, no animals were captured more than three
times; whether this indicated a larger home range or a
faster turnover (due to death or dispersal) was unknown.
There is little or no mortality of Podomys directly
attributable to fires on my study sites (Table 18), for two
possible reasons. Because of the uneven distribution of
litter and bare patches of sand, the effects of these fires
occur in a mosaic of varying intensity. These are not
catastrophic fires that burn or suffocate rodents, as oc¬
casionally observed in other habitats (Chew et al., 1959;
Tevis, 1956b). Secondly, mice are protected by remaining
within the tortoise burrows. Although logs, holes in trees,
and burned out root channels are used as temporary shelters,
as reported for Peromvscus by Baker (1974), Wolff and Hurl-
butt (1982), and others, I have no evidence that my animals
used long-term nests in logs or other structures above

120
TABLE 18. Survival of Podomvs following three prescribed
burns on the Ordway Preserve. N indicates the total number
of mice captured during the month prior to fire; D indicates
the percentage that disappeared during the month of the
fire; A indicates the percentage alive during the month
after the fire.
Site
N
D
A
BP
4
0
100
LP
3
0
100
SL
15
13
87

121
ground. Likewise, Layne (1970) considered Podomvs to be
exclusively burrow-dwelling.
I also compared trapping success on burned and unburned
sites by examining trapping effort expressed as trapnights
per total number of captures per hectare (Table 19); i.e.,
how much effort is required in an area before finding a
mouse? No effort was made to make trapnights equivalent,
but the total number of trapnights is listed for each year.
Regardless of the number of trapnights, more time was needed
to catch a mouse on the unburned sites. For all five years
on the SL sites, the numbers of trapnights required to catch
a mouse were much smaller and more consistent than on the
unburned areas.
What is the effect of prescribed burns on young ani¬
mals? From 1984 to 1988, I marked 17 young animals (juven¬
iles and subadults) on the unburned side of SL and 81 young
on the burn. I arbitrarily defined "resident" as an animal
that was trapped at SL for more than six months and found
that only 18 (10 females and 8 males) of those 98 young
animals became residents on SL (Table 20). A higher per¬
centage of the residents remained on the burned part of the
sandhill: 3% of the juveniles and 9% of the subadults re¬
mained on the unburned side, and 15% of the juveniles and
11% of the subadults remained on the burn.
The Podomvs population at SL reached its highest level
in 1984 (Fig. 13) and persisted at lower numbers through

122
TABLE 19. Comparison of trapping effort on unburned and
burned sandhills, where TN is the total number of trapnights
and trapping effort is expressed as trapnight/capture/ha
(TN/CAP/HA).
Unburned Site
Year
TN
TN/CAP/HA
BP
1988
830
19.53
LP
1987
371
51.53
SL
1984
476
7.39
1985
784
14.74
1986
786
280.71
1987
429
15.32
1988
704
8.38
Burned Site
Year
TN
TN/CAP/HA
BP
1988
1612
8.70
LP
1987
492
6.51
SL
1984
1444
0.64
1985
2308
1.00
1986
2958
1.17
1987
1839
0.99
1988
2789
1.02

123
TABLE 20. Residence of young Podomvs on the SL sandhill,
where N is the total number of young marked and R is the
percentage of marked young that became residents on that
sandhill.
N
R
BURNED
81
26
NOT BURNED
17
12

124
1988. The decline did not appear to be due to fire, but
might be related to food supply. For example, Kantola
(1986) reported that turkey oaks on the Preserve produced
many more acorns in 1983 than 1984. If acorns are a major
food of Podomvs. then failure of acorn crops might have a
negative effect on mouse populations, although feeding
experiments in high pine and sand pine scrub habitats have
yet to show a positive response to supplemental food (Young
and Stout, 1986 and Diet above). Populations at BP and LP
were consistently lower than at SL.
Responses of Peromvscus to fires in a variety of habi¬
tats are listed in Table 21. Beck and Vogl (1972), Tester
(1965), and Hedlund and Rickard (1981) noted that small
mammals can survive fires by remaining in burrows. Most
studies were short-term research on P. maniculatus. which
showed a positive response to fires in almost all habitats.
Population increases of Peromvscus have been correlated with
changes in food, particularly seeds and insects (e.g.,
Tevis, 1956a; Ahlgren, 1966; Layne, 1974; Krefting and
Ahlgren, 1974; Bock and Bock, 1983; Kaufman et al., 1983).
The only evidence of qualitative differences in diets comes
from Halford (1981), who showed that P. maniculatus on burns
consumed fewer Lepidoptera and more Coleóptera and plant
material than mice on unburned sites. Other authors related
fluctuations of Peromvscus populations to changes in litter
and other structural qualities of the environment (Tester

125
TABLE 21. Responses of Peromvscus species to fire. R =
response (I = increase, D = decrease, 0 = no response) and
Time is the duration of the study (in months).
State Habitat R Time
p.
californicus
CA
mixed brush
D
30
CA
chaparral
D
48
P.
aossvDinus
AL
longleaf pine
I
18
FL
slash & longleaf
I
36
LA
loblolly-shortleaf
I
~11
LA
loblolly-shortleaf
I
varied
plots
P.
leucoDus
KS
tallgrass prairie
0
24
WI
brush prairie
savanna
D
1
P.
maniculatus
AZ
ponderosa pine
I
~8
CA
grassland
I
48
CA
Douglas fir
Ib
7
CA
grassland
I
30
mixed brush
0
30
CA
mixed conifers
& hardwoods
I
?
Reference
Cook 1959
Lawrence 1966
Boyer 1964
Layne 1974a
Shadowen 1963
among
Hatchell 1964
Kaufman et al. 1983
Beck & Vogl 1972
Lowe et al. 1978
Lawrence 1966
Tevis 1956a
Cook 1959
McDonald 1983

126
TABLE 2l--continued
State
Habitat
R
Time
Reference
CO
lodgepole pine
I
3
Roppe & Hein 1978
ID
mixed conifers
I
1
Stout et al. 1971
ID
sagebrush
Ic
7
Halford 1981
KS
tallgrass prairie
I
24
Kaufman et al. 1983
Peterson et al.1985
MN
jack pine
I
36
Ahlgren 1966
MN
mixed conifers &
Krefting &
hardwoods
Id
144
Ahlgren 1974
MN
jack pine & aspen-
fir-spruce
D
2
Buech et al. 1977
MT
larch-Douglas fir
I
~60
Halvorson 1982
OH
oat monoculture
D
10
Crowner & Barrett
1979
OR
Douglas fir
I
132
Gashwiler 1970
OR
Douglas fir
I
21
Hooven & Black 1976
SD
ponderosa pine
& pine savanna
I
24
Bock & Bock 1983
WI
brush prairie
savanna
I
1
Beck & Vogl 1972
BRITISH COLUMBIA
mixed conifer
I
8
Sullivan 1980

127
TABLE 21—continued
State Habitat
R
Time
Reference
MANITOBA
jack pine
I
36
Sims & Buckner 1973
ONTARIO
black spruce &
mixed wood
I
~36
Martell 1984
P. oolionotus
AL longleaf
I
18
Boyer 1964
FL longleaf pine/
turkey oak
0
15
Arata 1959
GA old field
I
6
Odum et al. 1973
P. truei
CA mixed brush
D
30
Cook 1959
CA chaparral
D
48
Lawrence 1966
"Peromvscus sdd."
MN oak/savanna
(tallgrass prairie)
I
2
Tester 1965
PA mixed oak
De
15
Fala 1975
12 years later there was no significant difference in
numbers on burned and unburned sites.
b Numbers were lower on uncut areas.
Numbers increased after burn; there were, however, more
animals on the control sites.
d Then decreased after 7 years.
Numbers increased in the first month, then decreased.

128
1965; Beck and Vogl, 1972; Lowe et al., 1978; Blankenship,
1982). Martell (1984) reported that P. maniculatus on burns
had significantly fewer botfly larvae than mice on clear-
cuts. The success of P. maniculatus in particular on burned
areas has been attributed to its preference for open areas
and to increased food supplies (Peterson et al., 1985),
possibly combined with an increase in the number of shelters
provided by logs (Lowe et al., 1978). Competition with other
species might be a factor in some communities. Halvorson
(1982) studied P. maniculatus in larch-fir forest, where he
suggested that deer mouse populations were affected by
competition with Clethrionomvs qapperi. but mice gained an
advantage on open situations (e.g., burns) and in years when
conifers produced large seed crops.
When fires in the high pine are suppressed, wiregrass
might become more abundant but other herbaceous species
decrease (Laessle, 1942; Hilmon and Hughes, 1965). On the
Ordway Preserve, Gates and Tanner (1988) reported greater
plant diversity on burned sandhills than on unburned con¬
trols, with greatest diversity occurring on sandhills burned
within two years. At other sites, Hilmon and Hughes (1965)
found greatest diversity three years after fire. They also
suggested (1965:252) that "in central and southern Florida,
however, where some growth occurs year-round, herbage stays
green through the first winter after a fire; and sufficient
rough to carry another fire accumulates only after two

129
years." Abrahamson and Abrahamson (1989) demonstrated that
not all plants produce higher-quality fruit after fires even
if vigorous sprouting occurs. Not enough is known about the
diet of Podomvs to determine whether fire affects popula¬
tions of Florida mice through altering food supplies.
Conclusions
The prescribed burns on three high pine study sites on
the Ordway Preserve did not have major impacts on mortality
or the size and locations of home ranges of Podomvs. In
comparisons of burned and unburned sandhills, more trapping
effort was required to capture animals on the unburned
sites. Juveniles and subadults caught on the unburned sites
are apparently dispersers; very few remain as residents on
the unburned areas. Over a period of several years, numbers
on the unburned sites were less predictable and less stable
than on burned areas. On the unburned sites, I captured all
mice at peripheral burrows (on the edge of the unburned
sandhill near old pasture or burned sandhill habitats), and
in some years I found no animals. Additional experiments
with more replications (and burned and unburned sites of
equal area) are needed, but these might be difficult to
perform at Ordway, given the large home ranges and fluctua¬
ting populations of these animals.
The exact mechanism responsible for the effect of fire
on Podomvs was not determined. What is clear is that the

130
cessation of fire allows invasion of woody species into the
high pine and a subsequent conversion to xeric hardwood or
mixed pine forests (Laessle, 1958; Myers, 1985), in which
Podomvs are absent and species richness declines (Campbell
and Christman, 1982; Humphrey et al., 1985; Brand, 1987;
Gates and Tanner, 1988; Layne, in press). Although Peromvs-
cus and other small mammals are abundant after fires in
other habitats, it is tempting to speculate that the close
association of the gopher mouse and gopher tortoise con¬
tributes to the success of Podomys in the sandhills. In
fact, Gopherus and the species that utilize its burrows
benefit indirectly from fire (Auffenberg and Franz, 1982;
Jackson, 1985; Landers, 1987; Breininger et al., 1988).
Fires serve to maintain an open canopy and stimulate new
growth in plants that serve as forage for tortoises and
possibly facilitate movement by hatchlings (Cox, Inkley, and
Kautz, 1987 ) .
Finally, I wish to emphasize how little is known about
the effects of fires on small mammals. Before 1955, there
was little interest in the effects of fire on small mammals
(Taylor, 1981). Many of the early papers about fire and
small, non-sciurid rodents reflect the forester's interest
in rodents as predators of seeds and seedlings (e.g., Boyer,
1964; Ahlgren, 1966), and, as I have shown, many studies
were of short duration. It is still rather surprising,
however, that with the prevalence of both prescribed burns

131
and lightning strikes in the southeastern United States,
that there is still so little research on effects on the
non-game animals of the region. Between 1975 and 1979, only
two papers were published regarding effects of fire on small
mammals in the southeast (Taylor, 1981). Studies of long
duration that examine seasonal effects of fire, costs of
dispersal, and reactions of additional species are needed.

SUMMARY AND DISCUSSION
The purpose of this project was to examine the micro¬
habitat use by P. floridanus in the high pine. I attempted
to answer five questions:
1) Do Podomys in the high pine depend exclusively on
tortoise burrows?
2) How large are home ranges?
3) Do dietary data explain why some tortoise burrows
are preferred?
4) Is the local distribution of these mice predic¬
table?
5) Do populations respond positively to burns?
To answer these questions, I monitored one population
(SL) on the Ordway Preserve for five years. I also trapped
on three additional sandhills (AC, BP, LP) for shorter
periods of time and observed animals in a captive colony at
the University of Florida.
Field workers at Ordway (Eisenberg, 1983; Brand, 1987;
this study) have found Podomys to be generally restricted to
high pine and old fields, where they are the only murid
residents. Hammocks on the preserve are occupied by P.
qossypinus and Ochrotomvs nuttali, but Podomys might invade
the edges of xeric hammocks in drought years, if tortoise
burrows are available and if densities of the other murids
132

133
decrease. We suspect, but have not tested, that competitive
exclusion occurs in most years and that Podomvs is re¬
stricted to the high pine.
In my fieldwork I confirmed that, at least in the high
pine on Ordway, Podomvs reside exclusively in tortoise
burrows. In more than 33,000 trapnights I observed mice
entering holes of unknown origin only three times. Trap
success at burrows (mean = 7% on SL) is higher than on grids
(means = 0.5, 0.7, and 3.6% on three grids), as Eisenberg
(1983) suggested. Florida mice certainly climbed plants and
utilized fallen branches while foraging, but the typical
escape response to trapping (at both grids and burrows) was
to enter tortoise burrows. Mice modify burrows by con¬
structing chimneys and U-shaped tunnels that open into the
ceilings and sides of the main tortoise burrow. These
structures probably served as escape routes from predators
(presumably snakes) and they allowed occupation of burrows
after tortoises left and the main entrances collapsed. I
observed Podomvs and Rana areolata simultaneously utilizing
burrows, but interactions with other burrow residents are
unknown. Florida mice probably are both predators and prey
of other burrow commensals.
When trapping at burrows, I captured other burrow
residents, particularly R. areolata. Although G. volans
were captured at burrows 38 times (Table 1), there was no
evidence that they actually resided in burrows. Other

134
rodents (P. qossypinus, S. hispidus, 0. palustris) were
found on the sandhills rarely, although they are known to
utilize tortoise burrows (Jackson and Milstrey, 1989). The
absence of other small rodents might be an indication that
Ordway has harsher sandhills—more xeric with more unpredic¬
table rainfall and food supplies—than those of other study
sites. It would be interesting to compare population sizes
and physical attributes of high pine sites on a state-wide
basis.
At the SL burrows 255 individuals were marked between
August 1983 and November 1988, 32% of which were captured
only once. Mean trappability was 57%, with individual
trappability ranging from 14 to 100%. As reported by Layne
(1966) for populations in Alachua County, I found the major¬
ity of reproduction occurring between August and November.
Juveniles could be marked outside what I believed to be
natal burrows at approximately three weeks of age, judging
by pelage and weight (9-14 g). Both pre- and post-
reproductive dispersal appeared to occur, although I had no
means of determining mortality. On the SL sandhill, 8.6% of
all marked animals were present for 360 days or more; of
these animals, half of the females were first marked as
juveniles, whereas most of the males were marked as sub¬
adults or adults. The longevity records were 649 days for
females and 920 days for males.

135
Almost nothing is known about the diet of Podomvs, and
studies of fecal or stomach contents are badly needed. In
the field I observed mice eating insects, young shoots
(Smilax), and Asimina fruits. Captive animals accepted a
wide variety of acorns, pine seeds, and other fruits and
flowers collected on Ordway, and cached favored foods. My
preliminary tests of acorn preferences suggested that spe¬
cies other than Q. laevis might be preferred, but more
strenuous tests need to be performed. Like that of Young
and Stout (1986), my supplementation experiment did not have
a positive effect on wild populations, suggesting that
factors other than food determine local distributions of
Podomvs.
I examined home range size using three techniques:
fluorescent powder, mark-and-release trapping at grids, and
mark-and-release trapping at burrows. The first two techni¬
ques severely underestimated home range size. With the
burrow data, I obtained mean annual home ranges of 2601 m2
(SD = 1315.1) for 35 females and 4042 m2 (SD = 2269.4) for
40 males, using the Exclusive Boundary Strip Method
(Stickel, 1954; Davis, 1956). These means might be slightly
underestimated, since some of these adults utilized edges of
my study areas, but these figures were more realistic than
those obtained on 100 m2 grids. Home ranges of Podomvs have
not been reported previously, but based on the unpredic¬
tability of climate and food in the high pine, I would

136
predict home ranges in the scrub would be smaller. Adult
females had mutually exclusive home ranges; this observation
agreed with laboratory observations of litter mortality and
aggressive behavior when unrelated females were housed
together. Adult males had significantly larger home ranges,
which overlapped with ranges of other adults.
Local distributions of Podomvs were not solely depen¬
dent on numbers of tortoise burrows. For example, in their
research on the Citrus Tract (Citrus and Hernando counties),
Beckwith (1964) and Umber (1975) noted that Florida mice
usually were captured on sandhills with native vegetation,
rather than on sandhills converted to pine plantations,
although tortoise density appeared to increase on the plan¬
tations. During the course of this study mice practically
disappeared from AC, although plenty of tortoise burrows
were still present. Clearly, distributions of Podomvs cannot
be predicted from burrow density alone.
On two sandhills (BP in 1988 and SL in 1985, 1987, and
1988) I sampled the vegetation in circular quadrats centered
on 42 burrows to determine whether the presence of mice
could be predicted on the basis of local plant structure or
species composition. I found that the total number of cap¬
tures was correlated (p < 0.05) with stem densities of
turkey oak, longleaf pine, and persimmon, and with increased
numbers of herbaceous species. Minimum numbers of in¬
dividual mice could not be predicted as accurately. More

137
than 83% of the burrows sampled were reclassified correctly-
using the physical and vegetation parameters I measured
(Table 13); i.e., the parameters I used were useful predic¬
tors of the relative abundance of Podomvs at burrows.
Determining the response of Podomvs to prescribed burns
was more complicated than I expected. Five years of trap¬
ping might be too brief a period to determine population
responses, and so far, most studies of fire and small mam¬
mals have been of even shorter duration. When I corrected
for differences in area and trapping effort, I found that
apparent densities on the unburned sites of the BP, LP, and
SL sandhills frequently exceeded those on the burned sites.
However—and this can be seen particularly with the SL data
(Fig. 13)—apparent densities on unburned areas fluctuated
more wildly and frequently reached zero. I interpreted my
results as follows. In the short term, fires do not have a
negative effect on Podomvs, because I detected no mortality
or movements of home ranges attributable to burns. In the
long term, mice respond positively to fires, probably due to
the larger numbers of tortoise burrows, increased diversity
of herbaceous species, and the opening up of habitat. Total
numbers of captures of mice at BP and SL were correlated
with higher stem counts of longleaf pine (known to do best
on burned areas), higher stem counts of turkey oak and
persimmon (which produce extra shoots after fires), and
herbaceous diversity (which, according to Gates and Tanner

138
(1988), reaches its maximum on Ordway sandhills two years
after fire). On all of my study sites, we have not burned
frequently enough to eradicate adult turkey oaks and persim¬
mons, so these stem counts reflected coppicing due to fire
and not numbers of large trees.
Several suggestions regarding management of P. florida-
nus on Ordway can be drawn from this study. First, a con¬
tinuation of the present system of rotational burns (three-
year cycles, different seasons) on the sandhills appears
beneficial. Secondly, until we learn more of diet and
interspecific interactions, maintaining a mosaic of sand¬
hills and old fields would be a wise strategy. Conversion
of SL to longleaf pine savanna or BP to turkey oak savanna
might be detrimental to the species if diversity in plant
composition is lost. Thirdly, protection and assessment of
gopher tortoise populations should continue, under the
assumption that whatever benefits Gopherus populations will
benefit Podomvs and other burrow residents as long as diver¬
sity in the high pine vegetation also is protected.
Although Podomvs conserves water less well than Pero-
myscus and develops hypothermia at 10 C, it resides in
tortoise burrows on the high sandhills of the preserve where
Peromvscus does not. Podomvs seems to be a good colonist in
these areas and, as noted, its behavior differs from that of
scrub populations. Podomvs appears to be "k-selected" when
compared to Peromvscus. Mace and Eisenberg (1982) noted the

139
large brain size relative to head and body weight; they
associated relatively large brains with smaller litters,
diet and habitat complexity, and larger numbers of sympatric
congeners. This brain-body size relationship is consistent
with earlier suggestions that the present distribution of
Podomvs is relictual.

APPENDIX
This is a list of plant species (excluding grasses and
sedges) identified in vegetation samples around tortoise
burrows at Blue Pond (BP) and Smith Lake (SL). Each species,
and its frequency of occurrence in the high pine at Ordway
(unless another habitat is noted), was identified using
"Vascular Plants of the Ordway Preserve", compiled by David
W. Hall and Richard Franz. The taxonomic arrangement of this
list follows that of Wunderlin's "Guide to the vascular
plants of central Florida" (1982). Voucher specimens were
deposited in the Ordway Herbarium.
FAMILY PTERIDACEAE.
Pteridium aquilinum (L.) Kuhn. Bracken fern. SL. Abundant.
FAMILY PINACEAE.
Pinus clausa (Chapm. ex Engelm.) Vasey ex Sarg. Sand pine.
SL. Scattered individuals.
Pinus elliotti Engelm. Slash pine. BP.
Pinus palustris Mill. Longleaf pine. BP,SL. Characteristic.
Pinus taeda L. Loblolly pine. BP. Characteristic in loblol¬
ly pine-bottomland hardwoods and some old fields.
FAMILY ARECACEAE.
Serenoa repens (Bartr.) Small. Saw palmetto. BP,SL. Oc¬
casional .
FAMILY COMMELINACEAE.
Commelina erecta L. Spreading dayflower. SL. Common.
Cuthbertia qraminea Small. Roseling. BP,SL. Common.
Tradescantia roseolens Small. Scrub spiderwort. SL. Uncom¬
mon.
FAMILY SMILACACEAE.
Smilax auriculata Walt. Wild bamboo. BP,SL. Abundant in
xeric live oak forests and ruderal sites.
FAMILY AGAVACEAE.
Yucca spp. Haw. Weak-leaf yucca. BP,SL. Common.
FAMILY MYRICACEAE.
Mvrica cerifera L. Southern bayberry or wax myrtle. SL.
Abundant.
140

141
FAMILY FAGACEAE.
Quercus chapmanii Sarg. Chapman's oak. SL. Common.
Ouercus qeminata Small. Sand live oak. BP,SL.
Quercus incana Bartr. Bluejack oak. BP,SL. Common.
Quercus laevis Walt. Turkey oak. BP,SL. Characteristic.
Quercus hemisphaerica Bartr. Laurel oak. BP,SL. Abundant on
ruderal sites.
Quercus myrtifolia Willd. Myrtle oak. SL. Common in sand
live oak hammocks.
Quercus stellata Wang. Small post oak. BP. Uncommon.
Quercus sp. L. Possible hybrid. BP.
FAMILY ARISTOLOCHIACEAE.
Aristolochia serpentaria L. Snakeroot. BP.
FAMILY POLYGONACEAE.
Erioqonum tomentosum Michx. Dog's tongue or wild buckwheat.
BP,SL. Abundant.
Polyqonella gracilis (Nutt.) Meisn. Wireweed. BP,SL. Abun¬
dant .
FAMILY CARYOPHYLLACEAE.
Paronychia patula Shinners. Spreading whittlow-wort. BP,SL.
Stipulicida setacea Michx. Wireweed. BP. Common.
FAMILY RANUNCULACEAE.
Clematis reticulata Walt. Vase-vine. BP,SL. Uncommon.
FAMILY ANNONACEAE.
Asimina incarna (Bartr.) Exell. Flag pawpaw. BP,SL. Abun¬
dant .
Asimina pyqmaea (Bartr.) Dunal. Dwarf pawpaw. BP. Uncommon.
FAMILY CAPPARACEAE.
Polansia tenuifolia Torr. and Gray. Pineland catchfly. SL.
Common.
FAMILY ROSACEAE.
Prunus serótina Ehrh. Wild cherry. SL. Uncommon on ruderal
sites.
FAMILY CHRYSOBALANACEAE.
Licania michauxii Prance. Gopher-apple. SL. Abundant.
FAMILY FABACEAE.
Aeschynomene viscidula Michx. Prostrate jointvetch. BP,SL.
Common in old fields.
Astragalus obcordatus Ell. SL. Uncommon.
Cassia chamaecrista L. Partridge pea. SL. Common.
Chapmannia floridana Torr. and Gray. Alicia. BP,SL. Abun¬
dant .
Clitoria mariana L. Butterfly-pea. BP,SL. Uncommon.

142
Crotalaria rotundifolia (Walt.) Gmel. Rabbit-bells. BP,SL.
Common.
Palea pinnata (Walt, ex Gmel.) Barneby. Summer farewell.
BP. Common.
Desmodium strictum or D. tenuifolium Torr. and Gray. BP,SL.
Common.
Desmodium viridiflorum (L.) DC. Velvet-leaved tick-trefoil.
BP,SL. Common.
Ervthrina herbácea L. Cherokee bean. SL. Common in hammocks.
Galactia reqularis (L.) BSP. Prostrate milk-pea. BP,SL.
Common.
Lespedeza hirta (L.) Hornem. Hairy bush-clover. SL.
Psoralea canescens Michx. Buckroot. BP,SL. Uncommon.
Rhynehosia cinerea Nash. Ashy Rhynchosia. BP.
Rhvnchosia reniformis DC. Dollar-weed. BP,SL. Common.
Schrankia microphvlla (Dry. ex Smith) Macbr. Smooth-leaved
sensitive briar. BP,SL. Common.
Stylosanthes biflora (L.) BSP. Pencilflower. SL.
Tephrosia chrysophvlla Pursh. Golden hoary-pea. BP,SL.
Common.
Tephrosia florida (Dietr.) Wood. Long-stalked hoary-pea.
BP.
Tephrosia spicata? (Walt.) Torr. and Gray. Sand pea. SL.
Common in old fields.
Tephrosia virqiniana (L.) Pers. Goat's rue. BP,SL. Abun¬
dant .
FAMILY POLYGALACEAE.
Polyqala grandiflora Walt. Large-flowered polygala. SL.
Uncommon.
FAMILY KRAMERIACEAE.
Krameria lanceolata Torr. Sandbur. SL. Abundant.
FAMILY EUPHORBIACEAE.
Chamaesvce cordifolia (Ell.) Small. Round-leaved spurge.
BP.
Cnidoscolus stimulosus (Michx.) Engelm. and Gray. Bull-
nettle. BP,SL. Abundant.
Croton arqyranthemus Michx. Silver croton. BP,SL. Abundant.
Croton qlandulosus L. Tropic croton. SL.
Stillinqia sylvatica L. Queen's delight. BP,SL. Common.
Traqia urens L. Eastern tragia. BP,SL. Common on old
fields.
FAMILY EMPETRACEAE.
Ceratiola ericoides Michx. Florida rosemary. SL. Oc¬
casional .
FAMILY ANACARDIACEAE.
Rhus copallina L. Winged sumac. BP,SL. Abundant.

143
FAMILY VITACEAE.
Vitis aestivalis Michx. Summer grape. BP,SL. Common on old
home sites.
FAMILY HYPERICACEAE.
Hypericum hypericoides (L.) Crantz. St. Andrew's-cross. SL.
Common.
FAMILY CISTACEAE.
Lechea patula Legg. Narrow-leaved pinweed. SL.
Lechea sp. L. Pinweed. BP,SL.
FAMILY VIOLACEAE.
Viola septemloba LeConte. Seven-lobed violet. SL. Uncommon.
FAMILY TURNERACEAE.
Pirigueta caroliniana (Walt.) Urban. Smooth stem piriqueta.
BP,SL. Common.
FAMILY CACTACEAE.
Opuntia humifusa (Raf.) Raf. Prickly-pear cactus. BP,SL.
Common.
FAMILY APIACEAE.
Erynqium aromaticum Baldw. ex Ell. Fragrant eryngium.
BP,SL. Common.
FAMILY ERICACEAE.
Vaccinium arboreum Marsh. Sparkleberry. BP,SL. Occasional.
Vaccinium corymbosum L. Highbush blueberry. SL. Common in
mesic hardwood hammocks.
Vaccinium myrsinites Lam. Shiny blueberry. BP,SL. Common.
Vaccinium stamineum L. Deerberry. SL. Common.
FAMILY EBENACEAE.
Diospyros virginiana L. Persimmon. BP,SL. Abundant.
FAMILY APOCYNACEAE.
Amsonia ciliata Walt. Bluestar. BP,SL. Abundant.
FAMILY ASCLEPIADACEAE.
Asclepias humistrata Walt. Sandhill milkweed. BP,SL. Com¬
mon.
Asclepias tuberosa subsp. rolfsii (Britt.) Woods. Fid-
dleleaf butterfly-weed. BP. Uncommon.
Asclepias verticillata L. Whorled-leaf milkweed. BP,SL.
Uncommon.
FAMILY CONVOLVULACEAE.
Stylisma patens (Desr.) Myint. Trailing stylisma. BP,SL.
Common.

FAMILY POLEMONIACEAE.
Phlox nivalis Lodd. Trailing phlox. SL. Uncommon.
144
FAMILY VERBENACEAE.
Callicarpa americana L. American beauty-berry. SL. Common.
Stylodon carneus (Medic.) Mold. SL.
FAMILY LAMIACEAE.
Salvia azurea Lam. Blue sage. BP. Uncommon.
Scutellaria multicrlandulosa (Kearney) Small. Rolled-leaf
skullcap. SL. Common.
FAMILY SCROPHULARIACEAE.
Linaria floridana Chapm. Florida toadflax. SL. Common.
Seymeria pectinata Pursh. Sticky seymeria. BP,SL. Common.
FAMILY ACANTHACEAE.
Dyschoriste oblonqifolia (Michx.) Kuntze. Twin flower. SL.
Common.
FAMILY RUBIACEAE.
Galium pilosum Ait. Bedstraw. BP.
Hedvotis procumbens (Gmel.) Fosberg. Fairy footprints.
BP,SL. Common.
FAMILY ASTERACEAE.
Arnoolossum floridanum (A. Gray) H. Robins. Indian-plan-
tain. SL.
Aster tortifolius Michx. White-topped aster. BP,SL. Local¬
ly common.
Balduina anqustifolia (Pursh) Robins. Yellow buttons.
BP,SL. Common.
Berlandiera subacaulis (Nutt.) Nutt. Common greeneyes.
BP,SL. Abundant.
Carphephorus corvmbosus (Nutt.) Torr. and Gray. Large¬
headed carphephorus. BP,SL. Abundant.
Cirsium sp. Mill. SL.
Convza canadensis (L.) Cronq. Dwarf horseweed. BP.
Elephantopus elatus Bertol. Florida elephant's foot. BP,SL.
Abundant.
Eupatorium compositifolium Walt. Dog fennel. BP,SL. Abun¬
dant .
Eupatorium iucundum Greene. Ageratina. BP/SL. Abundant.
Euthamia tenuifolia (Michx.) Greene. Flat-topped goldenrod.
BP. Uncommon.
Helianthus anqustifolius L. Narrow-leaved sunflower. BP.
Hieracium meqacephalon Nash. Large-headed hawkweed. SL.
Common.
Hieracium sp. L. Hawkweed. SL.
Liatris pauciflora Pursh. Lopsided blazing-star. BP,SL.
Abundant.

145
Liatris tenuifolia Nutt. Fine-leaf blazing-star. BP,SL.
Abundant.
Phoebanthus grandiflora (Torr. and Gray) Blake. BP,SL.
Uncommon.
Pitvopsis graminifolia (Michx.) Nutt. Silk-grass. BP,SL.
Abundant.
Pterocaulon pycnostachvum (Michx.) Ell. Blackroot. SL.
Uncommon.
Pvrrhopappus carolinianus (Walt.) DC. False dandelion.
BP,SL. Uncommon.
Rudbeckia hirta L. Black-eyed susan. BP. Common on road¬
sides and old fields.
Silphium compositum Michx. Rosin weed. BP,SL. Common.
Solidago chapmanii Torr. and Gray. Chapman's goldenrod. BP.
Common.
Verbesina heterophvlla (Chapm.) Gray. Crownbeard. BP. Rare.

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BIOGRAPHICAL SKETCH
Cheri Jones was born in Albuquerque, New Mexico, on
December 1, 1957, one of two children to Charlene and Clyde
Jones. As the family of a mammalogist, the Joneses lived in
Louisiana, Virginia, Colorado, Texas, and Equatorial Guinea,
West Africa. At various times over the years her parents
maintained bats in the basement, hyraxes in the bathroom,
hamsters in the den, and monkeys in the family room. It is
hardly surprising that both offspring developed environmen¬
tal interests and scientific careers.
She earned her B.A. in 1979 from the Department of
Biology at Hastings College and her M.S. in biology in 1982
from Fort Hays State University. While attending the Uni¬
versity of Florida, she was a member of the Florida Commit¬
tee on Rare and Endangered Plants and Animals, spent a
semester in Costa Rica, and served as an officer for the
U.F. chapter of Graduate Assistants United. Her profes¬
sional affiliations include the American Association for the
Advancement of Science, American Institute of Biological
Sciences, American Society of Mammalogists, Florida Academy
of Sciences, Gopher Tortoise Council, North Dakota Natural
Science Society, Organization for Tropical Studies,
159

160
Southwestern Association of Naturalists, Sigma Xi, and the
Wildlife Society.

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.
\F. Eisenbercj^Chairman
Katharine Ordway Prhfegsor of
Ecosystem Conservations
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.
--
Jónn H. Kaufmann
Professor of Zoology
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.
(La fylíjÁ'
Richard A. Kiltie
Associate Professor of Zoology
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.
francis E. Putz
Associate Professor^of Botany

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.
Tohn G. Robinson
Associate Professor of Forest
Resources and Conservation
This dissertation was submitted to the Graduate Faculty
of the Department of Zoology in the College of Liberal Arts
and Sciences and to the Graduate School and was accepted as
partial fulfillment of the requirements for the degree of
Doctor of Philosophy.
May 1990
Dean, Graduate School

UNIVERSITY OF FLORIDA
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