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FLORIDA
MUSEUM OF

NATURAL HISTORY

ECOLOGY OF WHITE-TAILED DEER
IN EASTERN EVERGLADES NATIONAL PARK

AN OVERVIEW

Tommy R. Smith, Cynthia G. Hunter,
John F. Eisenberg, and Melvin E. Sunquist

Volume 39, No. 4 pp. 141-172 1996

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UNIVERSITY OF FLORIDA GAINESVILLE



Numbers of the BULLETIN OF THE FLORIDA MUSEUM OF NATURAL HISTORY are
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This journal is printed on recycled paper.

ISSN: 0071-6154 CODEN: BF 5BA5

Publication date: July 17,1996 Price: $1.50



ECOLOGY OF WHITE-TAILED DEER IN EASTERN
EVERGLADES NATIONAL PARK--AN OVERVIEW

Tommy & Smithi, Cynthia G. Hunter~,
John F. EisenbergJ, and Melvin E. Sunquist

ABSTRACT

A population of white-tailed deer (Odocoileus virginianus seminolus) was studied
in the eastern Everglades National Park from 1987 to 1992. The data for the interval
January 1987 to June 1991 are summarized in this publication. During this interval, from
35 to 40 deer were monitored annually. Deer were captured utilizing a shoulder-held gun
projecting a net, and fired from a helicopter. Captured deer were fitted with a radio collar
and, thereafter, located by telemetry from both the ground and a fixed wing aircraft.
Aerial locations were logged to a recognizable hectare square. Minimum convex polygons
of deer home range use then were determined. Individual females occupied ranges of
approximately 37 ha and males 113 ha, but seasonal differences in range use indicated v
that some males may have an annual range exceeding 12 km'. There was considerable -4
overlap of home ranges, especially among individual females and their presumptive
offspring. The population density was estimated to be 0.68 deer/km:. Adult females
produce a single young; twinning was not recorded. During the course of this study,
recruitment into the population was low. Mortality rates varied from year to year,
especially for young fawns. The population appeared to be stable. During the course of
the study, the panther (Puma coneolor) was present. Deer comprised 57 percent of 99
identified panther kills in the eastern Everglades National Park and adjacent areas. Adult

'T. R. Smith was f6rmerly a Wildlife Scientist at U.S.N.P., Evergla£les National Park. and now resides in
La Neuve, Belgium.

2C. G. Hunter is a 0-aduate student in Zoology at Witwatersrand University, South Africa
3 J, F. Eisenberg holds the Ordway Chair of Ecosystem Conservation at the Florida Museum ofNatural

History and School of Forest Resources and Conservation, University of Florida  Gainesville FL 32611-
780OUS.A

4 M. E. Sunquist is Associate Professor, Department of Wildlife Ecology and Conservation, University of
Florida, Gainesville FL 32611 U.S.A

Smith, T. R., C. G. Hunter, J. F. Eisenberg, and M. E. Sunquist 1996. Ecology of white-tailed deer in
eastern Everglades National Park-an overview. Bull. Florida Mus. Nat. Hist. 39(4):141-172.



142 BULLETIN FLORIDA FLORIDA MUSEUM NATURAL HISTORY VOL. 39(4)

bucks with a mean age of 5.5 years were the dominant age/sex class killed (47% of all
deer). We conclude that, although densities of both deer and panthers are low in the
eastern Everglades National Park, they were normal and near carrying capacity or stable.

RESUMEN

Se estudi6 una poblacian de ciervos de cola blanca (Odocodeus vi,yinianus
seminolus) entre 1987 y 1992 en la parte oriental del Parque Nacional Everglades. Esta
publicaci6n sumariza la informaci6n obtenida entre enero de 1987 y junio de 1991.
Durante este periodo se monitorearon 35 a 40 ciervos al afto. Los ciervos fueron
capturados usando una red projectada por un rifle desde un helic6ptero. A cada ciervo
capturado se le coloc6 un radio collar gracias al cual los animales pudieron ser localizados
usando telemetria desde tierra o desde una avioneta. Las localizaciones obtenidas desde
la avioneta fueron registradas en cuadriculados de una hectarea. Los ambitos de hogar de ~
los ciervos fueron entonces determinados usando poligonos convexos minimos. Hembras
y machos ocuparon dmbitos de hogar de 37 y 113 Ha, respectivamente. Variaciones
estacionales indicaron que algunos machos habrian tenido ambitos mayores que 12 kmi
Los Ambitos de hogar se sobreimpusieron considerablemente. Las hembras y sus
presuntos descendientes ocuparon rangos de hogar sobrepuestos en gran medida. La
densidad poblacional fue estimada en 0.68 ciervos/km: Las hembras adultas parieron
una sola cria no registr~ndose mellizos. El reclutamiento de la  poblaci6n fue bajo durante
este estudio. La tasa de mortalidad vari6 de afto a afto especialmente en los cervatillos
j6venes. La poblacian apareci6 estable. Pumas (Puma concolor) estuvieron presentes
durante el curso de este estudio. Un 57% de las 99 presas capturadas por puma e
identificadas en la parte oriental del Parque Nacional Everglades fueron ciervos. Ciervos
adultos macho de 5.5 afios en promedio fueron la clase de edad y sexo dominante (47% de
todos los ciervos). Nosotros concluimos que aunque la densidad de ciervos y pumas en la
parte oriental del Parque Nacional Everglades es baja, ambas poblaciones son normales y
se encuentran cerca de su capacidad de carga.

ACKNOWLEDGEMENTS

The principle investigators (JFE and MES) would like to thank the many
participants, sponsors, and advocates of this lengthy project, especially the National Park
Service and its employees. Sonny Bass' cooperation was essential during the fmal
preparation of this report.

The majority of the field work was organized and conducted by Tommy Rae Smith.
He deserves credit for continuing aerial locations of deer through many hardships.
Cynthia Hunter completed her master's thesis research on the reproduction of the white-
tailed deer in the Everglades National Park (ENP) and aided greatly in the preparation of
the final report. Jos6 Fragoso and Tadeu Oliviera served as part-time field assistants.
People in Homestead and ENP who were outstandingly helpful were so numerous that we
fear we may leave some unmentioned, but we must particularly thank Wyatt Enterprises'



SMITH ET AL.: WHITE-TAILED DEER IN ENP 143

pilots, Lynch Incorporated pilots, Sue Husari and the Fire Cache, John Ogden, Frank
Draughn, and W. B. Robertson.

Many thanks are due to the generous Geographical Information System (GIS)
computer specialists of the Department of Urban and Regional Planning at the University
of Florida, especially David Lambert, who took the initiative to make room for the 200
MB of data from the park so that Cynthia Hunter could sift through it and absorb some of
his good teaching.

This research was supported with funds from U. S. National Park Service Grants
5602519-12 and 5602497-12 under CA-5280-5-8004. Some supplemental funding
derived from the Katharine Ordway Chair of Ecosystem Conservation, University of
Florida, Gaines¥ille. The National Fish and Wildlife Foundation contributed monetarily
as well.

SCOPE AND HISTORY

As increasing attention, funds, and research efforts have been applied
to the understanding, restoration, and management of the Everglades
ecosystem, a proposal was approved to elucidate interactions between deer
(Odocoileus virginianus seminolus), Florida panthers (Puma concolor
coryi), and the vegetation, all thought to be unique and essential
components (Eisenberg and Sunquist 1986). The Florida panther, an
endangered subspecies, feeds primarily on deer and feral hog (Sus scrofa),
but the hog has been eliminated from National Park land; thus in Everglades
National Park (ENP), deer potentially provide the only large prey available
(Maehr et al. 1990; Alvarez 1993). However, there has been widespread
concern that the deer in the Everglades are at insufficient density and of
such small body size that they are possibly an inadequate source of
sustenance for the panthers (Belden et al. 1988). Information on deer
population dynamics was considered essential to management of the
panther. The Panther Recovery Plan (USFWS 1987) has been an important
ongoing aspect of park research, and this deer study was proposed and
approved to provide essential information.

Physiological, behavioral, and reproductive differences between the
race 0. v. seminolus and northern white-tailed deer have long been
suspected (Richter and Labisky 1985), but little was known of the ecology
of South Florida deer. Only one in-depth study of the natural history of this
subspecies had been conducted prior to the present research effort (Loveless
1959; Loveless and Ligas 1959) with a later study by Schemnitz (1974).

Deer in ENP are a sedentary (non-migratory) population. No hunting
has been allowed anywhere in ENP since its establishment in 1947, but
hunting continued uninterrupted in the forest reserves north of the park.



144 BULLETIN FLORIDA FLORIDA MUSEUM NATURAL HISTORY VOL. 39(4)

Mortality factors acting on deer in ENP were presumed to be panthers,
bobcats (Lynx ruBs), poachers, cars, and natural health factors, including
possibly nutrition, contaminated surface water, disease, and parasites. In
this regard we were interested in examining the relative impact of these
various mortality factors, the possible differential impacts on different age
classes of the population, and the implications for management when
community interactions were considered.

Originally a comparison of hunted and unhunted populations was
proposed (Eisenberg and Sunquist 1986). Richter and Labisky (1985)
found a significant difference in deer herd productivity on hunted vs.
unhunted sites of similar habitats within Florida. Generally, herd
productivity was higher on hunted sites, although densities, pregnancy rates,
fetuses/doe, and age structures varied between sites. Their study provided
strong impetus to study and manage the various sub-populations of deer and
panthers across Florida on a site-specific basis. This report summarizes a
study focused on the unhunted ENP population south and east of Shark
River Slough. A parallel study from 1989 to 1992 directed by R. F.
Labisky focused on the hunted populations in Big Cypress National
Preserve and adjacent deer in ENP north of Shark River Slough.

Deer density in the Everglades appeared lower than elsewhere in
Florida or the Piedmont (Harlow and Jones 1965; Newsom 1984), but the
Everglades deer had not been censused thoroughly. A survey-based census
of deer and panthers in ENP in 1970 (Schemnitz 1974) estimated 8
panthers and 1500 deer in the then 5261 km: ENP, yielding a density of
0.29 deer/lan:. Densities were even lower in the Water Management
Conservation Areas 2 and 3 north of the park at 0.10 deer/kn,2 (Schemnitz
1974). This difference was possibly due to more intensive hunting and
more severe flooding there (Loveless 1959). The only other authors to
survey 0. v. seminolus and as part of a state survey (Harlow and Jones
1965) found the Everglades ecosystem the one least able to support deer in
all of non-urban Florida. This was mainly believed to result from the low
nutrient availability for deer in the predominantly flooded prairie habitats.
Although seasonally flooded bottomland hardwood forests are thought of as
the best deer habitat in the Southeast (Sigler-Eisenberg and Eisenberg
1985), very little ofthis habitat exists in the Everglades (Fig. 1).

Since the early 1970s the Everglades deer herd has apparently declined
and remained at a very low density, relative to historical populations in the
Everglades (F. Dayhoff pers. comm. 1989). The only recorded large-scale



LAKE

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 ET AL.: W

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ITE

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D
 D

EER
 IN

 ENP 
145

1

11
]m
 . The original extent of the Evergla(les wetland.

0
The study site  centered on Taylor Slough. 2

0, with burm and road. 50 km.
..... Edges of aquatic flows,

Shad Slough & Taylor Slough.

L Big Cypress National Refuge

1 Everglades National Park's terrestrial portion.

Figure 1. Southern Florida and the now partitioned and partially drained Everglades National Park wetland.



146 BULLETIN FLORIDA FLORIDA MUSEUM NATURAL HISTORY VOL. 39(4)

die-offs were caused by tick eradication efforts in the 1930s, overhunting in
the conservation areas in the 1960s, and excessive flooding in 1957-1958
and in 1982-1983. In part the low densities were attributed to a lower
fecundity in the deer of the southern flooded region than even in the rest of
Florida (Harlow 1972), despite the state as a whole having deer of lower
fecundity than other deer herds north of Florida (Harlow and Jones, 1965).
These authors attributed the lower fecundity to possible mineral deficiencies
from the food plants growing on infertile soils (Deuver et al. 1986; Kushlan
1990). This study and the concurrent work on deer in the Big Cypress
National Preserve compared densities, reproductive output, predation, and
other mortality factors affecting deer in the Everglades.

STUDY SITE

The terrestrial habitats of ENP, covering 5660 km2, are mainly prairies
dotted with tree islands of various sizes and pine flatlands, The Everglades
is classified as 65% tropical savannah (USFWS 1979), but not much is
known regarding changes in this ecosystem's plant or animal components
since water control measures were implemented. The entire region can be
considered a hyperseasonal savannah (Sarmiento 1984), controlled by
annual flooding that used to coincide with the rainy season of May through
October, and now is managed by outputs from canals (Fig. 1) in a "rainfall
simulation strategy. Although this management scheme has produced wet
and dry seasons somewhat similar to the natural system (Walters et al.
1992; ENP unpubl. hydrology data), the volume of water has been slowly
decreased over the last 50 years, diminishing both high and low extremes of
depth that occur seasonally in the park at the southern end of the wetland
(Hunter 1990).

The Everglades ecosystem is believed to be stable in plant species
composition, due to the periodic disturbance of fire and water (Robertson
1953; Loveless 1959). The savannah's prairies are two types. Wet prairies
are dominated by one or more of the sedges sawgrass (Cladium
jamaicense), spikerush (Eleocharis sp.), and beakrush (Rhynchospora sp.).
These prairies are dotted with tree islands, or "hammocks," composed of
tropical hardwoods or "domes" of cypress (Taxodium disnchum). Dry
prairies are dominated by the grass called "muhley" (Muhlenbergia flipes)
and scattered with occasional or clumped hardwoods, especially willow
(Salix caroliniana), and where hydroperiod becomes longer, with the herbs
common to wet prairies. Parts of dry prairies have no hydroperiod (above



SMn'HET AL.: WHITE-TAILED DEER IN ENP 147

ground flooding) at all in some years. Scattered large areas of pinelands are
nearly monotypic (Pinus ellioni var. densa) forests with thick understories
dominated by Sabalpalmetto (Fig. 2).

The two areas ofthe park where water flows almost continuously year-
round are called "sloughs," and the study efforts of this report centered
around Taylor Slough. This site allowed access to the research facilities
and the "hole in the donut." This latter site is a disturbed area now invaded
by dense stands of exotic trees, such as Schinus terrebinth</bhus and
Melaleuca quinquenervia. More details of vegetation can be found in
Deuver et al. (1986) and Hunter (1990). Ongoing NPS studies of fire
management and regeneration, flooding regimes, and flowering phenology
were simultaneous with this study and provided useful habitat information
in the form ofunpublished data from Park Service personnel.

The climate is technically sub-tropical: Seasonality is dictated by
variation in rainfall, not temperature, although a frost occurs on an average
of once every ten years. Rainfall averages 148 cm per year, 81 percent of
which falls during the wet season, usually from late April through October
(Schomer and Drew 1982; Deuver et al. 1986). The intensity (* 34 cm per
year), timing, and duration of the wet season are highly variable beginning
April, May, or June and extending through August, September, or October.
Rainfall often occurs with two noticeable peaks, one in May or June and the
second in September (37 years ENP unpubl. hydrology data; Schomer and
Drew 1982). The effects of water on deer health and the synchronizing of
herd productivity were among the questions of interest addressed in this
study.

METHODOLOGY

General methods are described below, and more details concerning
methods are included in the Results section.

Capture

Initially, dartguns shot from blinds, buildings, and a truck were used in
an attempt to capture deer, but this method proved unsuccessful due to the
availability of thick cover for the deer. Box traps were not utilized. After
frequent visual sightings of deer in open prairies, the technique of utilizing a
shoulder-held gun projecting a net fired from a helicopter was attempted
(Barett et al. 1982; Chardonnet and Charity 1992). A gunner



148 BULLETIN FLORIDA FLORIDA MUSEUM NATURAL HISTORY VOL. 39(4)

D ~

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.

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• = depth gauge
, = depth, rainfall, and flow gauge

-·-· = abandoned, partially filled canals
= canal, with burm and road

---- = slightly elevated road only
P = pinelands
H = hardwood hammock
X = dense exotic shrubs

Figure 2. The study site in the Taylor Slough area of Everglades National Park, and the
prairies classified as either "wet" (W) or "dry" (D). "Intermediate" (I) prairies were wet
in 1988 and dry in 1989.



SMITH ET AL.: WHrrE-TAILED DEER IN ENP 149

experienced in the use of helicopter net-gun techniques was invited to
demonstrate, and 19 deer were caught in 15 hours of flight. This became
the chosen technique for capture and was subsequently used to obtain the
sample of deer collared, as represented in Table 1. Deer were also
recaptured using this technique.

Chase flights by helicopoter were limited in duration to a maximum of
about 10 minutes, varying slightly with environmental temperature.
Temperature and chase time were the key factors limiting capture-related
mortality, as long chases may result in capture myopathy. No chemical
immobilization was necessary: Deer were immediately tied, blindfolded,
freed from the net, and held on the coolest ground available for an average
of 20 minutes while measurements were taken and a radio collar affixed.
The pulse and respiration rates of captured deer were monitored while the
animal was restrained.

Data collected on each captured animal included: Date, time, location,
habitat, other animals nearby, total length, tail length, head length, ear
length, zygomatic width, neck circumference, height at shoulder, girth,
weight, hind foot length, front foot length, toe length, sex, tooth condition,
tail fat index, muscle index, pelage condition, ectoparasite index, and
samples of blood and vital signs taken. These data were collected by very
few personnel, so the subjective assessments were similar for each animal.

Stratification of Sample and Sampling Scheme

After the initial sample of 45 deer was obtained in 1987 by somewhat
random searching of open prairies around the HID area, deer could be
selected visually and not chased if of an age or sex that was not needed in
the sample. While not completely random nor completely representative of
the population, the capture sample represented fairly evenly those deer
available in open prairies.

Age was determined by tooth eruption and wear (Severinghaus 1949),
but with no previously collected jaws of known age from the Everglades,
accurate aging was only possible after the death of many deer: Jaws were
retained for laboratory analysis. Some deer near the park were shot to
obtain samples for assessment of physiological condition. These data are
compiled in reports to FGFC (see Forrester 1990, 1993b; McCowan 1988).

Radio-tagged deer were located by telemetry on a rotating diel
schedule that allowed one to three locations/animal/week, evenly spaced to
cover 24 hours. A subset of the sample was located more intensively for



150 BULLET[N FLORIDA FLORIDA MUSEUM NATURAL HISTORY VOL. 39(4)

the "fawning ecology study" (Hunter 1990). That study, conducted from
January through July of 1989, included the creation of a digital map of 106
microhabitat types over the Taylor Slough portion of the Park. This map
included water coverage, with the extent of surface water determined by
fying systematic transects and noting the sun reflecting from the water and
by walking to newly flooded zones and measuring water depths. This
information was used for analysis of behavior and habitat use by does
during the fawning season (Hunter 1990).

Telemetry and Mapping of Locations

Collared animals were located by either fixed-wing aircraft or a van.
The plane (Cessna 182) had two H-element antennae, one mounted below
on each wing strut. Antenna were connected to a receiver through a switch-
box which allowed signal reception to be checked on either side of the plane
as the pilot was instructed to home-in on the deer sought. The van was
equipped with an 2.4-m telescoping mast supporting two H-element
antennae set at the same height but 1.5 m apart. This mast could rotate
360° on a marked compass rose, and the two antennae were connected by a
null combiner, yielding a silent spot thus indicating best direction of signal
(Cochran 1980). When fully extended the antenna reached 29 ft above the
ground. Signals were clear up to 4 km from the target animals in this flat,
open terrain.

Aerial locations were 16gged on an orthophoto map of the site to a
recognizable hectare square by circling over the animal until landmarks
were recognized. Hence, this technique could only be used during daylight
hours. From dusk through dawn, locations were made from the van by
triangulation using two fixes on the same animal made within a 30-minute
span, or a shorter period if the animal was moving. Approximately 60% of
all locations were evenly represented throughout the diel and seasonal
cycles. The maps used for the population study were USGS orthophoto
maps on 7.5 minute quad sheets at a scale of 1:24,000, drawn in 1972 from
aerial photos. The fawning ecology study (Hunter 1990) used a detailed
microhabitat map created from 1986 aerial photos and digitized on the
Geographical Information System (GIS) to allow more detailed habitat
studies. The maps used for analysis of habitat use thus far include Florida
Game and Fish's 16-habitat map of Dade County, a 5-habitat broad scale
map created by the GIS lab at ENP, and a 176-microhabitat patch map of
Taylor Slough.



SMITH ET AL.  WHITE-TAILED DEER IN ENP 151

The program TELEM88 (Coleman and Jones 1988) was used to
delineate minimum convex polygons of deer home-range use. These
polygons were digitized for some study animals and overlaid on the various
vegetation maps, on which we had already quantified "availability" of
various habitats by area. "Use" was then compared to random and to
available proportions.

Questions relating to deer habitat use were: What are the major
habitats preferred by each age/sex group? Do deer utilize all habitats in
proportion to their availability? Do adult females change habitat use at
time of fawning for fawn-rearing? Do deer avoid or respond to rising
floodwaters: To what degree do deer and panther home ranges overlap?
What effects do fires have on deer, during and in the successional stages
following a fire? Do human land use practices affect deer? Can
management affect mortality factors of deer?

Other data collected during telemetric locations were activity, habitat,
and, if possible, associations with other deer (visual or simultaneous
reception). Each radio collar was equipped with an activity switch, which
was sensitive to movement of the collar, and activity or inactivity was
expressed by different signal pulse rates. Inactivity was triggered by the
collar being immobile for 60 seconds, and any motion of the head would
switch it back to the "active" phase. Signals were monitored for at least
three minutes per fix. If the animal was recorded as inactive for this entire
period, it was checked again several hours later. If an inactive signal was
obtained two or three times over a 24-hour period, the animal was assumed
to be dead or the collar had fallen off, and a walk-in was performed (defined
below). All visual locations of deer included a description of behavioral
activities and any associations, with great attempts to minimize or prevent
disturbance to the animals. For example, although most deer became
accustomed to the fixed-wing plane, they would sometimes look up at it,
thus triggering an active signal even though they were lying down or
resting. Ifthis was seen the animal was recorded as inactive. Most activity
readings were obtained before the plane was within a mile of the animal,
presumably before it would respond to the presence ofthe plane.

Population Censusing

Walk-ins were actual locating of an animal by the investigator on foot.
As mentioned above, this was routinely done if the animal was believed to
be dead, or in some cases, for visual observation of doe and fawn behavior.



152 BULLETIN FLORIDA FLORIDA MUSEUM NATURAL HISTORY VOL. 39(4)

Doe and fawn associations were recorded most often from three observation
towers placed in the home ranges of does (Hunter 1990). If a carcass was
found, the cause of death was assessed, and if possible, the lower jaw was
retrieved for aging. Simultaneously, kills made by collared panthers were
examined after the cat left and some jaws of uncollared deer were obtained
in this way.

Each radio location (fix) was evaluated for accuracy and only those
fixes with 51 ha error polygons (Heezen and Tester 1967) were used in the
analyses. Radio-collared deer were located biweekly in a stratified pattern
of locations spaced through the 24 hours and evenly distributed over the
sex/age categories of the sample. In addition, during the fawning seasons of
1988 and 1989, one ofus (Hunter) located a subsample of pregnant females
an additional 2-3 times per week to obtain an estimate of the fawning date.

Aerial censuses of the 425 km2 study area were flown systematically
approximately every other month, and two observers counted all marked
and unmarked deer seen along the flight path.

RESULTS AND COMPARISONS

Sample Size

Table 1 summarizes the captures and recaptures conducted each year.
Capture-related mortality was very low. The percent of all deer handled
that received injuries or died within three weeks of capture was 6 percent in
1988 and 3 percent in 1989 and 1990.

Sex and Age Structure of Population

As described in Methods, efforts were made to obtain a sample
representative of the population as it exists in the prairies of southeastern
ENP (Table 1). Aging of young deer was usually determined by the tooth
eruption schedule of Severinghaus (1949), whose sample was from central
Florida. The assumption that eruption would be the same in the Everglades
deer was not entirely true. In Severinghaus' study, the permanent third
lower premolar was completely erupted by 20 months, while in ENP many
yearlings known to be older than this retained the juvenile condition of a
deciduous lower premolar. Eruption was not complete in 63% of capture



SMITH ET AL.: WHITE-TAILED DEER IN ENP 153

Table 1. Captures by netgun 6f deer in eastern Everglades National Park.

Captures Recaptures Monitored

M FM M FM M FM

1987-1989
F 11 11 1 1 1 1
J231636
A 13 24 4 3 8 16

1989-1990
F721 144
J003 234
A 1 1 4 17 8 18

M= male; EM = female.F=<1 yr, J= 1-2 yrs; A=3+yrs.

animals 18-25 months of age, yet was complete in at least one 24-month
old, thus determination of 2-year-old status was not absolute using this
tooth-eruption method. We could not rely on tooth wear to accurately age
adults; it seems likely that the diet of deer in the Everglades is unique, and
the wear characteristics for each year are unrecorded. We began a
collection of known-age jaws to allow for this comparison in the future.
The most precise method of aging dead deer was from cementum annuli
counts. Of 64 jaws aged accurately by cementum annuli, the following age
groups were obtained. This sample was from collared animals and other
carcasses in the park. The ages identified are included in Table 2.

The pregnancy rate was determined by two methods, with slightly
different results. By visual examination of females from the air and at
capture, no fawns or yearlings appeared pregnant, and only 25 percent of
all yearling females were seen with fawns. However, a physiological
examination of does (Formster 1990, 1992b) from a sample collected
slightly to the north of ENP showed that 3 out of 3 yearlings were pregnant



154 BULLETIN FLORIDA FLORIDA MUSEUM NATURAL HISTORY VOL. 39(4)

Table 2. Age classes identified.

Years

Year 0-1 24 3-5 5-9 9-14 14+

1987-91 15 23 7 11 7 5

when killed, and from a sample of does greater than 2 years of age, 50%
and 79% of those collected in 1988 and 1989, respectively, were pregnant
when killed. This sample provides a minimum estimate of reproduction,
because some of the does were killed in October, during the rut, negating
the possibility oftheir fertilization just before or after the harvest.

Other estimated ages of live-captured animals indicated a stable age
distribution throughout the period of this study.

While physical size was not used to determine age, it was recorded
with an estimate of age and analyzed to assess the hypothesis that
Everglades deer are smaller than northern deer. Based on captured animals,
fawns weighed from 4 to 25 kg, yearlings from 24 to 34 kg, adult bucks
from 38 to 61 kg, and adult does from 26 to 43 kg. Other measurements
taken on captured animals are summarized in Table 3.

In the eastern ENP, for each year the percentages of collared and
monitored "adult" females observed with a live fawn were: 1987,67% (but
observation not through full fawning season--82% pregnancy detected);
1988, 86% (94% pregnancy detected); 1989, 94% (87% pregnancy
detected); and 1990 (91% pregnancy detected). None of the harvested
females carried twin fetuses, and only one sighting of a doe with twins
occurred during the entire study. Hence, twinning in the ENP appears to be
extremely rare, with survival of both young even rarer, thus gross
productivity is equal to the pregnancy rate, which we estimate to be about
90% for females 2 3 years and about 80% for 2-year-old females.

A fat estimate was taken on each animal handled using a scale of 1 (no
fat) to 6 (thick fat) by palpitating just above the base of the tail. Adult



SMITH ET AL.: WHITE-TAILED DEER IN ENP 155

Table 3. Physical characteristics of deer captured in eastern Everglades National Park
during 1989.

Weight' Total length2 Girth Hind foot~

SexAge N X Range X Range X Range X Range

4 Z k Z

22 23* 34.9 26-43 159.7 145-171 74.2 64-80 42.7 39-45
22 5 49.3 38-61 174.1 167-181 83.2 75-92 45.4 45-47

1 3' 26.3 25-30 149.3 143-153 67.3 62-71 40.8 41-42
1 3 30.8 29-34 155.3 150-161 71.0 67-76 43.3 43-45

I kilograms
2 total length, cm
3 length, cm
4 includes 4 deer collected in health study by Don Forrester
5 included 1 deer collected in health study by Don Forrester

males changed from a pre-rut mean fat index of 5 to a post-rut mean of 2.3.
Lactation had a similar effect on females, changing their pre-lactation (late
pregnancy) mean of 4 to a post-fawning mean of 1.3. This subjective index
is based on a small sample and it has limitations, but it does indicate a
seasonal loss of condition for both sexes, and a recuperation period
sufficient to enter the following breeding season in restored physical
condition.

Estimates of Population Size

Census flights along systematic serpentine transects were flown
approximately every other month, with two observers counting marked and
unmarked animals visible below and to the sides of the plane. This
technique yielded such highly variable results that no reliable density
estimate was obtained until the standard deviation of mean estimates shrank
to a reasonable level as the number of censuses increased. The mean
estimate of density over the 425 km: study area by December 1990 was
0.67 i 0.24 deer/knr (Table 4). While we cannot completely explain the



156 BULLETIN FLORIDA FLORIDA MUSEUM NATURAL HISTORY VOL. 39(4)

Table 4. Population characteristics estimated each year in eastern Everglades National
Park (also see Table 6).

1987 1988 1989 1990

N 35 35-40 -
M-A x 0.14 - -
M-J x 0.60 - -
M-F 0.67 0.36 - -
FR 0.78 0.80 0.71 1.0 (burned)

0.9 (unburned)
M:F 3:1 3.4:1 1:1 -
PopEst x 230 298 -
Density/lan: <1.0 0.55 0.67 0.68

N = number collared deer monitored for most of thatyear
M-A = adult mortality recorded among collared animals
M-F = fawn mortality recorded among collared or visually known fawns
M-J = juvenile mortality recorded among collared 1 -2 year olds
FR = rate of fawning in does 2 2 years old and ;suming no twinning
M:F = sex ratio ofobserved adults averaged over year
PopEst- estimate oftotal number ofdeer resident in the study area, 425 larn'
x = not estimated

variation found between surveys, some of it may be due to seasonal
movements of males (see reproduction), and the overall estimate is
comparable to those ofprevious authors (Loveless 1959; Schemnitz 1974).

Mortalily and Predation

Assessment of All Mortality Factors

Ectoparasite loads were minor to non-existent on ENP deer. For
example, only 4 of 44 deer captured in 1989 were found with detectable
ticks or mites, and these were low loads in September-November. A
veterinary survey of a random sample of deer from ENP and Big Cypress
National Park (BCNP) showed the populations to be in good health with
low parasite loads (McCowan 1990; Forrester 1992b). Parasites are not
believed to be a significant detriment to deer in ENP.



SMITH ET AL.: WHITE-TAILED DEER IN ENP 157

Age-related Mortality

Fawn mortality was significantly higher in wet prairies than in dry
prairies (Chiz < 0.01) (Hunter 1990); and it was greater in male fawns than
in females (Smith 199lb). While not all causes of fawn mortality could be
determined, the possible differences between the two prairie types are
unlikely to derive from different predator pressure. More likely the
mortality differences are related to vegetative/phenological differences in the
maternal treatment of male vs female fawns, if (for example) the female
offers more maternal protection to female offspring. The annual mortality
rate of yearlings was approximately 28%. For 2- to 3-year olds the rate
was approximately 16%, and for adults, 4-years old and greater, it was
about 22% per year. Rates were not significantly different according to sex
or age class, except for fawns as noted above.

A Leslie matrix analysis based on the estimated mortality rates
indicates that at least 60 percent survivorship (to adult reproduction) of
female fawns is necessary to maintain numbers of females in the population,
and hence provide stability. Survivorship of fawns into the "follower
stage," at about three months, is not uniform by sex or location: fawns of
both sexes died during this period in wet prairies (Hunter 1990). However,
survivorship of female fawns appears sufficient to allow population growth.
Observed population growth during the five years of study was lower than
the mortality data would predict. There is some (inconclusive) evidence
that a skewed sex ratio at birth may exist in the Everglades population,
favoring males.

Panther Predation and Comments on the Bobcat

The summaries in Table 5 indicate that adult deer are the primary prey
of resident panthers in South Florida, especially in ENP where hogs, the
other prey capable of sustaining adult panthers (Maehr et al. 1990; Alvarez
1993), are almost never found. Panther predation may be a significant
factor on deer populations where deer are hunted (Dalrymple and Bass in
press; Smith and Bass 1994). Where deer are hunted, e.g. BCNP, younger
deer are killed more frequently and bobcat predation on fawns seems to be
an important population factor (Boulay 1992; Sargent 1992). It should be
noted that panthers will kill bobcats, and where panthers are present, bobcat
nunnbers may be depressed.



158 BULLETIN FLORIDA FLORIDA MUSEUM NATURAL HISTORY VOL. 39(4)

Table 5. Predation of deer by collared panthers in Everglades National Park and vicinity.

No. panthers No. adult study No. study
monitored deer killed fawns

killed

1987 6 0 0*
1988 7 4 1
1989 7 6 1?
1990 8 9 4

* incomplete year

The panther population utilizing ENP remained small throughout the
study, but was thought to be at or near carrying capacity for the region
(Smith and Bass 1994). During the wet season, panthers reduced the area
in which they hunted and killed less deer from May through September
(Dalrymple and Bass in press). Deer comprised 57 percent of 99 identified
kills and 78 percent of the biomass killed by panthers in ENP 1987-1990
(Dalrymple and Bass in press). Adult bucks with a mean age of 5.5 years
were the dominant age/sex class killed (47% ofall deer killed).

Although mortality rates varied from year to year, especially among
young fawns, the overall rate of recruitment and mortality were consistently
low compared to other deer populations in South Florida. This implies that
the population is stable. Perhaps the low densities of deer and panthers
reflects limitation by environmental carrying capacity and by density
dependent £actors.

In a larger sample of fawns in the Everglades/Big Cypress border
region, mortality was similar among sexes. Male fawns had a greater
survival rite in only one of three years. Fawn survival rate was higher in
BCNP than in ENP (0.39 vs 0.10). Deaths were of higher frequency
between two and six months of age. Bobcat (Lynx mfis) predation
accounted for 58 percent of all deaths. Recruitment ranged from 11% to
26% in BCNP and 0-13% in ENP (Boulay 1992). Overall fawn
survivorship in that study averaged 24%, and loss was attributed mainly to
bobcat predation. The situation is not completely comparable to the



SMNH ET AL.: WHITE-TAILED DEER IN ENP 159

southeastern ENP, where panther predation exerts a greater impact, but the
biology of deer reproduction should be similar.

Reproduction

Males associated with females from July to October and with each
other in sporadic small groups the rest of the year. Males showed antler
velvet from March to mid-June and shed antlers in December and January.
Fawns were dropped from January through early May, with peaks in
February and March, the late dry season (Hunter 1990). Females seem to
associate in permanent matriarchal groups, although two does maintained
solitary home ranges throughout the study.

The overall pregnancy rate among females 2 2 years old at fawning
was 79 percent. This is a very conservative estimate, as we were unable to
verify fawning until the fawn was visible, which precludes any estimates of
early neonatal mortality. We can conclude that the females in ENP have a
high conception rate and sufficient males are available for mating. Yearling
does do not often seem to conceive. This could result from the sedentary,
matriarchal social system precluding the youngest females from breeding
access to males.

No twinning was observed in the study. Although only six does were
killed and examined for fetuses (Forrester 1992b), the careful monitoring of
a total of 22 does during the fawning season allows us to conclude that
twinning is very rare in the ENP, or if twinning does occur, the mothers are
unable to maintain two fawns, and one sibling usually dies soon after birth.

Fawning proved to be synchronized with the late dry season (Fig. 3)
(Hunter 1990; Boulay 1992) and was significantly later in does resident in
wet prairies vs those resident in dry prairies (Hunter 1990) (Fig. 4). This
was contrary to hypothesized differences in timing, where in wet prairies
fawn mortality due to high-water conditions might possibly constrain
reproductive success and thereby select for earlier fawning. Instead,
although there was significantly more fawn mortality in wet than in dry
prairies, fawning must occur later in wet prairies due to nutritional
constraints on does (Hunter 1990) (see also Fig. 5).



160 BULLETIN FLORIDA FLORIDA MUSEUM NATURAL HISTORY VOL. 39(4)

O.v.seminolus

cm. rain proportion fawns born
50 - - 0.8

Z-1

40-
- 0.6

30 -

-0.4

20 -

1 -0.2

10- HriuO 3,Vt!11111, N# * *,tr]*~:r.- I:"0
J FMAMJ.J ASOND J FMAMJ J ASOND J FMAM JJA

| 1987 | 1988 | 1989 |

- Rain MI Fawning ~• Mean Conception Date

wet & dry prairies combined
Figure 3. Fawning seasons in relation to rainfall averaged over the study site, Taylor
Slough region of Everglades National Park. Deer conceptions (by backdating from
fawning dates) occurred just before the peak of two very different rainy seasons (Hunter
1990).

# fawns born

1

Jan A Jan B Feb A Feb B Mar A Mar B Apr A Apr B
half-months

~ Dry Prairie (n-9) I Wet Prairie (n-9)

Figure 4. Number of fawns born in the two prairies types, 1988-1989, Taylor Slough portion of
Everglades National Park. Each block represents fawning events with median date in either the first 15
days ofthe month (A) or the rest ofthe month (B). The mean birthdate was significantly different between
the two prairies types (Hunter 1990).



SMITH ET AL.: WHITE-TAILED DEER IN ENP 161

60-

50
-0- DRY PRAIRIE -0- WET PRAIRIE

40

ce
nt
im
et
er
s

30

20

10

0
JFMAMJJASONDJFMAMJJASONDJFMAMJJA

1987 1 1988 1 1989

Figure 5. Rainfall on Taylor Slough's dry prairies (station R3110) and wet prairies
(station NP67) during this fawning study, Everglades National Park (Hunter 1990).

Home Range and Habitat Use--Results and Discussion

A serious constraint on the analyses of habitat use was the lack of
digital data on habitat availability. Although the park has a history of
extensive botanical work, and most of the area's flora are quite complete,
the quantification of coverage by each community could be assessed
accurately over such a large area only with the advent of digital remote
sensing data. ENP purchased in 1986 (available in 1987) a complete set of
false color-infrared tinted stereoscopic photographs of the entire
landholding. An initiation of digitizing and ground-truthing the habitat
information from a 10,000-scale resource provided the Taylor Slough
Microhabitat map (Hunter 1990). This map was used for a find-scale (106
microhabitat patch-types, 625 m2 cells) analysis of females' behavior
during the fawning season. The Florida Game and Fresh Water Fish
Commission meanwhile created a broad-scale habitat map (16 categories,



162 BULLETIN FLORIDA FLORIDA MUSEUM NATURAL HISTORY VOL. 39(4)

Table 6. Population characteristics for white-tailed deer in eastern Everglades National
Park for the interval January 1987-June 1991 (3 years). *

Average or RangeJan 87-Jun 91

Density (#Rm2) 0.68
Productivity (F/AF) 0.82-0.94
Sex ratio

at birth 1:0.7
at 1 yr or over 1:2.4

Litter size (n = 85) 1.0
Age structure (%)

0-1 yr 22
2-3 yrs 33
4-5 yrs 10
< 6 yrs 33

Mortality
Fawns 0.54
Yearlings 0.24-0.32
Adults (>2 yrs)

Females 0.13
Males 0.19

Recruitment per year (%)
(218 mon 53 yrs) 1

* Based in part on 35-40 collared deer monitored per year as well as observations of known-age deer.

1000 m2 cells) for the entire state, which was not completely ground-
truthed. The inaccuracies were corrected by Hunter after ground4ruthing
the previous map. The broad-scale map was used for a study of
the annual home ranges of males and females in the core study area,
centered around Long Pine Key (LPK). Also, some satellite imagery was
obtained from the Florida Department of Transportation and classified into
six broad landscape types for panther home-range analysis (Smith and Bass
1994).

Home-range size estimates for deer were calculated from telemetry
locations using the program TELEM88 (Coleman and Jones 1988), and the
core activity area was defined as the 80 percent isopleth. Panthers home



SMITH ET AL.  WHITE-TAILED DEER IN ENP 163

ranges for were estimated from telemetry locations by using the program
HOMERANGE (Samuel et al. 1985); the core activity area was defined as
the 60 percent harmonic mean.

Home ranges of adult male deer were larger than those of adult
females. Overlapping home ranges for 22 females covered 8.15 kmi for an
average individual area of 37 ha. However, taking overlap into
consideration, average core home-range area (80% isopleth) for females
was 430 ha. Home ranges (which barely overlapped) around the LPK area
for 16 adult males covered 18.05 knf, for an average individual area of
1.13 kin: (113 ha). Individual core areas for males averaged 12.4 kmZ
(1240 ha). Thus, the density of females was nearly four times higher in
these prairies than that of males, and they utilized more of the same habitats
and even ranges than males did. Females showed strong home range fidelity
throughout the study, exhibiting a matriarchal hierarchy of social
dominance with overlapping home ranges of related females. This pattern
also was seen in the BCNP deer populations (Zultowsky 1992). Zultowsky
(1992) found female home ranges averaging 5.4 kin2 in BCNP and 2.95 kmZ
in ENP. Again water levels did not restrict movement, but home ranges
expanded during the dry season. Sargent (1992) found home ranges of
males to be 7,0 lan: in BCNP and 2.9 lan: in ENP,

Adult bucks utilized dense cover year round much more than adult
females did. Collared females were rarely found in the large, dense stands
of pinelands, Schinus, or hardwoo(is, while most males' home ranges
included these stands. All deer seem to avoid the dense Schinus stands
present in the "Hole-in-the-Donut," but the location as well as capture of
deer in any dense tree cover was nearly impossible. Hence, captures
emphasized prairie residents, and initial conclusions that deer densities were
inversely proportional to distance from LPK or any large tree island (Smith
and Bass 1994) were compromised. Panthers and adult bucks did utilize
the edges of tree islands more than any other habitat (Smith and Bass
1994). This "edge effect" probably reflects a stronger need by males for
flight cover. Many does were resident year-round in open prairies with little
tree cover available. Wet season flooding did not result in any noticeable
shifts in home range use of deer (Hunter 1990). However, panther home
ranges were smaller in the wet season (Dalrymple and Bass in press), and
the cats avoided flooded areas, thus the increased fawn mortality in wet
prairies during the wet season cannot be explained by panther predation.

The severe drought of 1989 allowed the normal lightning strike-
induced fires to burn more widely than usual, and about half the study area



164 BULLETIN FLORIDA FLORIDA MUSEUM NATURAL HISTORY VOL. 39(4)

was burned in the "Ingraham Fire," which was a spectacle second only to
the Yellowstone blaze in recent NPS history. But since a policy of
controlled and natural burns had been maintained for many years preceding
the fire, it mainly cleared undergrowth and did not often kill mature trees.
The prairie vegetation showed regrowth with in a week, while the hardwood
hammocks and some pine lands took months to "green up" (ENP Fire
Management pers. comm.). Following the fire, deer that were resident in
the area did not deviate from normal patterns of home range use. The
natural periodic cycles of fire and flood have been (and still are) believed to
keep the vegetative communities of the Everglades at equilibrium over time
(Robertson 1953; Wade et al. 1980). The noted changes are attributed to
human influences on the wetland ecosystem (Walters et al. 1992). In the
year following the Ingraham Fire, deer were monitored inside and outside of
the burned zone. There was no notable migration of deer into or out of the
burned area at any time. Nor was there any notable change in home range
use during rising waters or when home ranges were 100 percent inundated
(as in the wet year of 1988) (Hunter 1990). However, historical maximum
depths were not reached, so a critical depth where water begins to affect
deer mobility was not determined. Harlow's (1959) suggestion that 30 cm
was the critical depth might be valid, but these depths rarely occur south of
the Water Conservation Areas (40 years ENP hydrology data) (Fig. 6).

The microhabitat study (Hunter 1990) analyzed only the fawning
females' use of 106 microhabitat patch types in wet and dry prairies,
determining their use versus availability. A principle-components analysis
is undenvay, indicating so far that shelter elements within the prairies, such
as tall grass and willow heads and other shrubbery, are important to does
with fawns. Also, the evidence supports the theory of Loveless (1959) and
Harlow (1961) that "lilies" (four species of aquatic herbs with flowers) are
important to the diets of deer, especially in wet prairies.

While investigating the relative rates of success in fawn-rearing in the
two types of prairie (wet or dry, defined above) (Hunter 1990; 1994), it was
discovered that does residing year-round in wet prairies had significantly
less success raising fawns to the 3-month-old follower stage, and that they
fawned on average one month later than those does living in the dry prairies.
These results suggested a possible nutritional dependency on the part of the
mothers during the early wet season, constraining successful lactation to
that period when blooming ofnutritious flowers occurs.

Miller (1993) contrasted home ranges of males and females with
respect to habitat type. In his thesis he distinguished two habitat categories:



SMITH ET AL.: WHrrE-TAILED DEER IN ENP 165

60- -

30-

60- -

57 68 69 6 es .62 83 64 65 66 67

60-

5 30-

10

S 30-
60-«»f«

67 68 69 70 71 72 73 74 76 76 17

60- -

30- A - --

30 - ~ -
00-

.........

77 78 79 80 81 82 83 84 85 86 87

years

Figure 6. Modelled water table in relation to ground level (horizontal line at 0), from data
recorded each year from 1957 through 1987, in typical wet prairie (station P38),
Everglades National Park. Year marks are at January (Hunter 1990).

(1) ridges with trees, and (2) wet prairies with scattered trees. Both males
and females had smaller home ranges on ridges (5.3 km2 and 2.5 km2) and
larger home ranges if inhabiting prairies (8.2 and 6.2 ). The ridges are
characterized by greater cover density and frequency of tree islands in
BCNP (Miller 1993).



166 BULLETIN FLORIDA FLORIDA MUSEUM NATURAL HISTORY VOL. 39(4)

GENERAL DISCUSSION

We found little seasonal variation in density and reproductive output of
this relatively K-selected subspecies. Although densities of both deer and
panthers are low in eastern ENP compared to ranges farther north, we
suspect that they are normal and near carrying capacity. The home ranges
and stable, matriarchal social system exhibited by the deer indicate that the
Evergla£les provides a meta-stable environment. Although primary
production is low, there is a seasonal peak as the floodwaters arrive, which
appears to be the factor influencing the onset of deer reproduction and
ultimately the successful rearing of fawns. Hence, reproduction is
synchronized, and mortality, although exhibiting seasonal fluxes, probably
is important, perhaps regulating the juvenile age class number.

Harlow (1961) conducted a thorough analysis of the nutrients available
to deer over most of Florida. In the Everglades, he quantified the nutrient
content ofthe 41 food items found in deer rumens, but unfortunately did not
assess availability. Although concentrations of minerals and inorganic ions
are generally lower in Taylor Slough than in the larger Shark River Slough
to the northwest (Schomer and Drew 1982), they were believed to be
sufficient in soil and. plant matter such that,minerals were not the limiting
factor to deer (Harlow 1961). Instead, the lack of cover and of edible
browse were believed to be limiting, at least in the wet prairies (Harlow and
Jones 1965). Camp (1932) reported that cows could not survive in wet
prairies of South Florida during flooded months, but their larger body size
(and energetic advantage) allows them to obtain a nutritionally better diet
than that on which deer can survive in the same habitat. Comparing deer
diets in the Everglades to those in the rest of Florida, Harlow and Jones
(1965) claimed the prairies to be the habitat least able to support high
densities of deer. The current study showed that deer can and do survive
solely in prairie habitats at low densities and in good health. Densities
found in this study were very comparable to those of white-tailed deer in the
Venezuelan 11anos (Brokx 1972) (optimum 4/ km2 = 25 ha/deer; crude
density 2.0-2.5/lan2), a similar habitat, implying this is the carrying
capacity of flooded savannahs (Eisenberg et al. 1979; Stuwe 1985).
Authors have labelled bottomland hardwood forests as the best habitat for
deer in the southeastern United States (Sigler-Eisenberg and Eisenberg
1985), but the hardwood forests in the Everglades are scattered fragments
utilized for cover and browse, especially by bucks, and do not characterize
the terrain in terms of dominance. It is possible that the hardwood tree



SMITH ET AL.: WHITE-TAILED DEER IN ENP 167

islands are the "best" habitat for deer and are monopolized by the adult
males, as has been noted in more crowded deer populations at other locales
(LaGory et al. 1991). But it is unlikely that at these low densities, deer of
either sex in ENP would be excluded from any habitats by density-
dependent factors. Perhaps the more solitary social system of the males
promotes utilization of tree cover, while the more grouped females need
open habitats to promote cohesion.

The vegetative cover consists of structural characteristics ofgrasslands
as well as interspersion of shrubs and trees, higher interspersion providing
better habitat for deer (Moore and Haas 1973). It seems likely that there is
sufficient cover for female deer and fawns in the prairies with minimal tree
island stands and that the presupposed dependency on tree islands (Loveless
1959; Miller 1992) is not necessarily a selective force shaping behavior of
Everglades deer.

Flooding did not appear to be a direct mortality factor in the typical
Everglades. However, it may be a limiting stress factor in dry prairies,
especially to lactating does. This anecdotal evidence supports the theory
(McNab 1963) that animals in xeric regions will have larger home ranges
than their conspecifics in mesic regions, which was supported by
Rautenstrauch and Krausman (1989) with mule deer (0. hemionus crooki).
The difference in apparent carrying capacity of prairies in the wet season,
and the timing of fawning, is apparently not due directly to the hazards of
water, but more to its effects on the vegetation.

Fire management of Everglades habitats did not prove beneficial to
deer. Unlike some northern hardwood forests, which are maintained and
encouraged to regenerate by fire, pine flatwoods with palmetto understories
are affected by fire, and open wet prairies need to be burned more than they
are to hold back hardwood encroachment, but this encroachment might be
beneficial to the deer. A better land management strategy for the sake of
deer and panthers would be to acquire and protect from disturbance land in
areas which, if disturbed, have a potentially high susceptibility to
floodwater or saltwater intrusion. The entire Everglades ecosystem is in a
state of transition due to pollution, water diversions, urban encroachment,
and now Hurricane Andrew. The restoration efforts should utilize the
information presented here on the previous stable status of deer and
pantliers as a minimum baseline. While it is possible the Park can only
support 6-8 panthers (the number that has been used since at least the
1960s [Schemnitz 19741), it should not be assumed that the ecosystem was



168 BULLETIN FLORIDA FLORIDA MUSEUM NATURAL HISTORY VOL. 39(4)

supporting its maximum capacity of deer, and there is good evidence that
deer might have been the factor limiting panthers (Smith and Bass 1994).

NOTE: This publication is an abbreviated version of the project final repon submitted to the U.S.N.P. by J.F.E. and
ME.S. The priority of authorship reflects the actual field efforts and data analyses. The total data set for ENP,
developed by T.R. Smith, will be analyzed and submitted for separate publications by Smith and associates.

lfE.

LITERATURE CITED AND REFERENCES

Alvarez, K. 1993. Twilight of the panther. Myakka River Publishing, Sarasota FL, 501
P.

Barett, M. W., J. W. Nolan, and L. D. Roy. 1982. Evaluation of hand-held net gun to
capture large mammals. Wildl. Soc. Bull. 10:108-114.

Belden, R. C., W. B. Frankenberger, and S. T. Schwikert. 1988. Panther habitat use in
south Florida. J. Wildl. Mgmt. 52(4):660-663.

Blouch, R. A. 1987. Reproductive seasonality of the white-tailed deer on the Colombian
11anos. Pp. 339-343 in C. M. Wemmer, ed. Biology and management of the
Cervidae. Smithsonian Press, Washington, D. C.

Boulay, M. C. 1992. Mortality and recruitment of white-tailed deer fawns in the wet
prairie/tree island habitat of the Everglades. M.S. thesis, Univ. Florida,
Gainesville, 77 p.

Branan, W. V.,and R. L. Marchinton. 1987. Reproductive ecology of white-tailed and
red brocket deer in Suriname. Pp. 344-351 in C. M. Wemmer, ed. Biology and
management of the Cervidae. Smithsonian Press, Washington, D. C.

Brisbin, I. L., and M. S. Lenarz. 1984. Morphological comparisons of insular and
mainland populations of southeastern white-tailed deer. J. Mamm. 65(1):44-50.

Brokx, P. A. 1972. A study of the biology of Venezuelan white-tailed deer (Odocoileus
vignianus gnnotis Wiegmann, 1883), with a hypothesis on the origin of South
American cervids. Ph.D. dissertation, Univ. Waterloo, Ontario, Canada.
. 1984. South America. Pp. 525-546 in L. K. Halls, ed. The white-tailed deer:
Ecology and Management. Stackpole Press, Harrisburg PA.

Bunnell, F. L. 1987. Reproductive tactics of Cervidae and their relationships to habitat.
Pp. 145-167 in C. M. Wemmer, ed. Biology and management of the Cervidae.
Smithsonian Press, Washington, D. C.

Byers, C. R., R. K. Steinhorst, and P. R  Krausman. 1984. Classification of a technique
for analysis of utilization-availability data. J. Wildl. Mgmt. 48:1050-1053.

Camp, P. D. 1932. A study of range cattle management in Alachua County, Florida.
Florida Agric. Exp. Sta. Bull. 249,28 p.

Caughley, G. 1977. Analysis of vertebrate populations. John Wiley and Sons, New
York.



SMITH ET AL.  WHITE-TAILED DEER IN ENP 169

Chardonnet, P., and S. Charity. 1992. A marsh deer (Blastocerus dichotomus) capture
experiment in Brazil using a net-gun from a helicopter. Pp. 583-588 in F. Spitz, G.
Janeau, G. Gonzalez, and S. Aulagnier, eds. Ongults/Ungulates 91. Proc. Internatl.
Symp., Toulouse, France, September 2-6, 1991. S.F.E.P.M.-I.R.G.M., Paris-
Toulouse, France.

Cochran, W. W. 1980. Wildlife telemetry. Pp. 507-520 in S. D. Schemnitz, ed. Wildlife
management techniques manual, 4th Ed. Wildl. Soc., Washington, D. C.

Coleman, J. S., and A. B. Jones, III. 1988. User's guide to TELEM88: Computer
analysis system for radio-telementry data. Res. Ser. 1, Dept. Fisheries Wildl.,
Virginia Polytec. Inst. State Univ., Blacksburg, 49 p.

Dalrymple, G. H., and O. L. Bass. In press. Food habits of the Florida panther in
Everglades National park, Florida. Bull. Florida Mus. Nat. Hist. Vol. 39.

Deuver, M. J., J. E. Carlson, J. F. Meeder, L. C. Deuver, L. H. Gunderson, L. A. Riopelle,
T. R. Alexander, R. L. Myers, and D. P. Spangler. 1986. The Big Cypress National
Preserve. Nat. Aud. Soc. Res. Reprint 8, Key Largo.

Downing, R. L., B. S. McGinnes, R. L. Petcher, and J. L. Sandt. 1969. Seasonal changes
in movements of white-tailed deer. In L. K. Halls, ed. Proceedings of a symposium
on white-tailed deer in the southern forest habitat. U. S. Southern Forest Exp. Sta.

Eisenberg, J. F., and M. E. Sunquist. 1986. Ecology of the white-tailed deer in
Everglades National Park. Proposal to N.P.S.

Eisenberg, J. F., M. A. O'Connell, and P. V. August. 1979. Density, productivity, and
distribution of mammals in two Venezuelan habitats. Pp. 187-210 in J. F.
Eisenberg, ed. Vertebrate ecology in the northern neotropics. Smithsonian Inst.,
Washington, D. C.

Forrester, D. J. 1990. Studies on the health and diseases of white-tailed deer in
Everglades National Park and Big Cypress National Preserve: Annual progress
report. Coop. Agreement CA-5000-7-8007, Dept. Infectious Diseases, Coll. Vet.
Med., Univ. Florida, Gainesville, 10 p. + figs.
. 1992a. Florida panthers and bobcats. Pp. 174-203 in Parasites and Diseases of
Wild Mammals in Florida. Univ. Presses Florida, Gainesville, 460 p.

1992b. White-tailed deer. Pp. 275-353 in Parasites and Diseases of Wild
Mammals in Florida. Univ. Presses Florida  Gainesville, 460 p.

Harlow, R. F. 1959. An evaluation of white-tailed deer habitat in Florida. Tech. Bull. 5,
Florida Game and Fresh Water Fish Comm., Tallahassee, 69 p.

1961. Fall and winter foods of Florida white-tailed deer. Quart. J. Florida Acad.
Sci. 24(1):19-38.
. 1972. Reproductive rates in white-tailed deer in Florida. Quart. J. Florida Acad.
Sci. 35(4):165-170.
, and F. K. Jones, eds. 1965. The white-tailed deer in Florida. Tech. Bull. 9,
Florida Game and Fresh Water Fish Comm., Tallahassee, 240 p.

Harmoning, A. K. 1976. White-tailed deer dispersion and habitat utilization in central
North Dakota. North Dakota Game and Fish Dept., P-R W-67-R 13, 14, 15 Phase C
Rept. 199-A.

Haugen, A. 0., and D. W. Speake. 1957. Parturition and early reactions of white-tailed
deer fawns. J. Mamm. 38:420-421.

Heezen, H. R., and J. R. Tester. 1967. Evaluation of radio-tracking by triangulation with
special reference to deer movements. J. Wildl. Mgmt. 31:124-141.



170 BULLETIN FLORIDA FLORIDA MUSEUM NATURAL HISTORY VOL. 39(4)

Heisey, D. M., and T. L. Fuller. 1985. Evaluation of survival and cause specific
mortality rates using telemetry data. J. Wildl. Mgmt. 49:668-674.

Hirth, D. H. 1977. Social behavior of white-tailed deer in relation to habitat. Wildl.
Mono. 53,55 p.

Hirst, S. M. 1975. Ungulate-habitat relationships in a South African woodland/savanna
ecosystem. Wildl. Mono. 44,60 p.

Huegel, C. N. 1985. Predator avoidance behaviors that favor fawn survival in white-
tailed deer in south-central Iowa. Ph.D. dissertation, Iowa State Univ., Ames,
150 p.

Hunter, C. G. 1990. Odocoileus virginianus seminolus: The ecology of fawning in wet
and dry prairies. M.S. thesis, Univ. Florida, Gainesville, 138 p.

1994. Effects of prairie hydroperiod on timing and success of fawning in
Everglades deer. Unpubl. Rept. Dept. Wildlife & Range Sciences, Univ. Florida,
Gainesville.
, R. Lemos de sa, and T. Oliviera. 1989. A GIS analysis of Taylor Slough,
Everglades National Park. Unpubl. Rept., Dept. Wildl. Range Sci., Univ. Florida,
Gainesville.

Kushlan, J. A. 1990. Freshwater marshes. Pp. 324-363 in R. L. Myers and J. J. Ewel,
eds. Ecosystems of Florida. Univ. Central Florida Press, Orlando.

LaGory, K. E.,C  Bagshaw III, and I. Lehr Brisbin II. 1991. Niche differences between
male and female white-tailed deer on Ossabaw Island, Georgia. Appl. Anim. Beh.
Sci. 29:205-214.

Loveless, C. M. 1959. The Everglades deer herd: Life history and management. Florida
Game & Fresh Water Fish Comm. Tech. Bull. 6.
, and F. J. Ligas. 1959. Range conditions, life history and food habits of the
Everglades deer herd. Trans. No. Amer. Wildl. Conf. 24:201-214.

Maehr, D. S., R. C. Belden, E. D. Land, and L. Wilkins. 1990. Food habits of panthers
in southwest Florida. J. Wildl. Mgmt. 54:420423.

McCowan, J. W. 1988. Big Cypress watershed deer/hog/panther relationships E-1-12
animal performance report. Florida Game & Fresh Water Fish Comm., Tallahassee.

McNab, B. K. 1963. Bioenergetics and the determination of home range size. Amer.
Nat. 97:130-140.

Miller, K. E. 1993. Habitat use by white-tailed deer in the Everglades: Tree islands in a
seasonally flooded landscape. M.S. thesis, Univ. Florida, Gainesville, 105 p.

Moore, K. R., and R. H. Haas. 1973. Remote detection of deer habitat factors. NASA
Tech. Rept. RSC-50, Remote Sensing Ctr., Texas A&M Univ., College Station, 113

P.
Newsom, J. D. 1984. The coastal plain. Pp. 367-380 in L. K. Halls, ed. White-tailed

deer: Ecology and Management. Stackpole Press, Harrisburg PA.
Nue, C. W., C. R. Byers, and J. M. Peek. 1974. A technique for analysis of utilization-

availability data. J. Wildl. Mgmt  38:541-544.
Rautenstrauch, K. R., and P. R. Krausman. 1989. Influence of water availability and

rainfall on m ovements of desert mule deer. J. Mamm. 70(1):197-201.
Richter, A. R., and R. F. Labisky. 1985. Reproductive dynamics among disjunct white-

tailed deer herds in Florida. J. Wildl. Mgmt  49:964-971.
Robertson, W. B. 1953. A survey of the effects of fire in Everglades National Park.

Unpubl. Rept., Natl. Park Serv., Homestead, 169 p.



SMITH ET AL,: WHITE-TAILED DEER IN ENP 171

Sarmiento, G. 1984. The ecology of neotropical savannas. Harvard Univ. Press,
Cambridge MA.

Samuel, M. D., D. J. Pierce, E. 0. Garton, L. J. Nelson, and K. R. Dixon. 1985. User's
manual for program HOMERANGE. Tech. Rept. 15, Forestry, Wildl. Rang Exper.
Sta., Univ. Idaho, Moscow, 70 p.

Sargent, R. A.,Jr. 1992. Movement ecology of adult male white-tailed deer in hunted
and non-hunted populations in the wet prairie of the Everglades. M.S. thesis, Univ.
Florida, Gainesville, 66 p.

Schemnitz, S. D. 1974. Populations of bear, panther, alligator and deer in the Florida
Everglades. Florida Sci. 37(3):157-167.

Schomer, N. S., and R. D. Drew. 1982. An ecological characterization of the lower
Everglades, Florida Bay, and the Florida Keys. FWS/OBS-82/58.1, Office Biol.
Serv., U. S. Fish & Wildl. Serv., Washington DC, 246 p.

Seber, G. A. F. 1982. The estimation of animal abundance. 2nd Ed. Charles Griffin &
Co.,London, 654 p.

Severinghaus, C. W. 1949. Tooth development and wear as criteria of age in white-tailed
deer. J. Wildl. Mgmt. 13:195-216.

Sigler-Eisenberg, B., and J. F. Eisenberg. 1985. Variation in white-tailed deer
populations: A note on the southeast coastal plain. Ann. Mtg. S. E. Archeol.
Assoc., Pensacola FL.

Smith, T. R. 1991. Recruitment and loss in an unhunted subtropical white-tailed deer
population. P. 176 in R. D. Brown, ed. The biology of deer. Springer-Verlag, New
York.

1991. Effects of hydrology and vegetation on the distribution and abundance of
white-tailed deer and Florida panthers in Everglades National Park. Unpubl. Doc.,
Ann. Rept. FY 1991, South Florida Res. Ctr.„ Homestead, 26 p.
, and O. R. Bass. 1994. Landscape, white-tailed deer, and · the distribution of
Florida panthers in the Everglades. Pp. 693-708 in S. M. Davis and J. C. Ogden,
eds. The Everglades: Ecosystem and its Restoration. St. Lucie Press, Key Largo,
Florida.
, J. F. Eisenberg, and M. E. Sunquist. 1989. Ecology of white-tailed deer in
Everglades National Park (CA-5280-5-8004): Progress January 1987-January 1989.
Rept. to S. E. Reg. Off. Natl. Park Serv., Atlanta.

, and . 1990. Ecology of the white-tailed deer in Everglades National
Park. Natl. Fish & Wildl. Found. Proj. 89-68, Final Rept

, and . 1990. Ecology of white-tailed deer in Everglades National
]Park(CA-5286-5-8604): Progress February 1989-January 1990. Rept. To S. E.
Reg. Off. Natl. Park Serv., Atlanta.

Stuwe, M. 1985. Aspects of structure and reproduction of white-tailed deer populations,
Odocoileus virginianus, in Venezuela and Virginia. Saugetierkundliche
Mitteilungen 32:137-141.

U. S. Fish and Wildlife Service. 1987. Florida panther (Felis concolor coryi) recovery
plan. Prepared by Florida Panther Interagency Committee, Atlanta, 75 p

Wade, D., J. Ewel, and R. Hofstetter. 1980. Fire in south Florida ecosystems. U. S.
Dept. Agric., Forest Serv. Gen. Tech. Rept. SE-17.

Walters, C., L. H Gunderson, and C. S. Holling. 1992. Experimental policies for water
management in the Everglades. Ecol. Applic. 2(2):189-202.



172 BULLETIN FLORIDA FLORIDA MUSEUM NATURAL HISTORY VOL. 39(4)

Wemmer, C., ed. 1987. Biology and management of the Cervidae. Smithsonian Inst.
Press, Washington DC.

Zultowsky, J. M. 1992. Behavioral and spatial ecology of female white-tailed deer in the
Everglades ecosystem. M. S. thesis, Univ. Florida, Gainesville, 81 p.



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