of Management Amphibians,

advertisement
This file was created by scanning the printed publication.
Errors identified by the software have been corrected;
however, some errors may remain.
Management of Amphibians,
Reptiles, and Small Mammals
in Xeric Pinelands of
Peninsular Florida1
I. Jack S t ~ u tDonald
,~
R. Ri~hardson,~
and
Richard E. Roberts4
Xeric pinelands seem incongruent
with reference to Florida, a state with
annual rainfall that ranges from 50-65
in (19.6-25.6 cm). None theless, the
Florida peninsula contains thousands
of acres of sandy soil derived from
marine deposits dating to the Pleistocene (White 1970).Two distinct plant
associations, longleaf pine (Pinus
palustris)/ turkey oak (Quercus laevis)
sandhill and sand pine scrub (Pinus
clausa), have developed on these nutrient deficient and excessively welldrained soils. Significant areas of
these plant associations occur at
higher, albeit modest, elevations relative to the surrounding landscape. In
fact, certain topographic features,
e.g., the Lake Wales Ridge and the
Marion Upland, were likely to have
been true islands during interglacial
periods while the remainder of Florida was covered by a shallow sea.
Regardless of their exact origin, xeric
pinelands support many relatively
unusual species of amphibians, reptiles, and small mammals.
'Paper presented at symposium, Management of Amphibians, Reptiles, and
Small Mammals in North America. (Flagstaff, AZ. July 79-21, 1988).
21. Jack Stout is Professor of Biological
Sciences, Department of Biological Sciences, University of Central Florida, Box
250CO, Orlando, FL 328 16-0368.
3DonaldR. Richardson is Adjunct Professor, Department of Biology, University of
South Florida, Tampa, FL 33620.
4RichardE. Roberfs is Biologist, Division of
Recreation and Parks, Florida Department
of Natural Resources, Hobe Sound, FL
33455.
Abstract.-The primary xeric pinelands of peninsular Florida are lon~leafpinelturkey oak sandhills and
sand pine scrub, Their management on public lands
is largely confined to prescribed burning to maintain
fire climax status of the vegetation. The regulation of
large-scale developments on private land has stim ulated interest in preserve design and management.
The suite of techniques used to solve conflict between natural system preservation and development includes: (I) conservation set asides (preserves) on site; (2) habitat restoration; (3) purchase
and dedication of off-site preserves; (4) species relocation; and (5) wildlife resource mitigation fund.
Human population growth (3.3%
per year) and development in Florida
continues to encroach on upland
habitats and particularly on xeric pinelands. Most of the habitat loss is to
agricultural uses, principally citrus.
Oddly, the state's excellent wetlands
protection acts have forced development into the uplands. Thus, xeric
pinelands and their narrowly
adapted fauna and flora are increasingly threatened by area reduction,
fragmentation and isolation.
It is our intent to discuss the management of these xeric pinelands in
general and, more specifically, in the
context of small preserves in an otherwise developed landscape. Management of xeric pinelands as ecosystems is yet in its infancy, and more
detailed prescriptions for designated
species are unproven. However,
progress is being made (Cox et al.
1987) and improvement and revision
of current thinking on management
practices is anticipated. This paper
summarizes selected literature on
xeric pineland and the species associated with these communities to assess management practices. In addition, unpublished information has
been used and identified in the text.
Preserve design efforts by us have
been on behalf of developers responding to development orders prepared by governmental agencies.
These designs are site specific in detail, but nonetheless point to general
problems and solutions. Our approach has been to focus on provid-
ing the area required to support
minimum viable populations of
"keystone" or otherwise critical animal species of a given xeric pineland.
Once this area is settled on, management should focus on those species
whose minimum area requirements
are met, whereas no special efforts
are expended on species with larger
area requirements.
XERIC PINELAND HABITATS
Longleaf Pinepurkey Oak
Sandhills
The longleaf pine/ turkey oak
sandhill association (LLP/TO) was
about 15% (2,110,256 ha) of the
natural landscape of peninsular Florida in pre-Columbian times (fig. 1)
(Auffenberg and Franz 1982).This
xeric pineland occupies rolling topography of several ridge systems
that run north-south, notably Trail
Ridge, the Lake Wales Ridge, and the
Brooksville Ridge; numerous lesser
ridges and hills are identified by
White (1970). These ridges consist of
deep, well-drained soils of the
Lakeland, Eustis, and Blanton associations (Beckenbach and Hammett
1962).Laessle (1942,1958a) describes
the LLP/TO plant association as a
fire climax system dominated by
longleaf pine; slash pine (Pinus elliotti) replaces longleaf pine in the
community in south Florida. Turkey
oak is a minor tree, but can achieve
co-dominance when fires are suppressed. The predominant understory plant is wiregrass, Aristida
stricta; however, a rich assemblage of
perennial herbs vary in prominence
in concert with seasonal changes.
Monk (1968) recognizes two additional phases of sandhill vegetation
in north central Florida: (I) longleaf
pine/sand post oak (Quercus nrargaretta) and (2) longleaf pine/southern
red oak (Quercus falcata). A fourth
phase, longleaf pine/scrub hickory
(Cay a floridam), occurs in the southern portion of the Lake Wales Ridge
(Abrahamson et al. 1984).Veno
(1976), Givens et al. (1984), and Abrahamson et al. (1984) provide quantitative data on LLP/TO community
structure and dynamics. Myers
(1985)suggests that longleaf pine/
turkey oak and sand pine scrub associations are successionally linked in
some portions of their geographic
ranges. Differences in physical /
chemical features of soils of LLP/TO
and SPS communities in the Ocala
National Forest are not considered to
be sufficient to explain the local dis-
Figure 1 .-Potential geographic distribution
of longleaf pine/turkey oak sandhill and
sand pine scrub xeric pinelands in Florida.
Light shading indicates the sandhills and
darker shading indicates the scrub. These
distributions are based on Davis (1 980) and
do not reflect minor sites of either community due to the scale of the itlustration.
tribu tion of the communities (Kalisz
and Stone 1984).
Prior to settlement by European
man, ground fires occurred in LLP/
TO sandhills at intervals of 1-5 years.
These relatively "cool" fires favor
regeneration of longleaf pine, flowering by grasses and herbs, and suppress growth of woody plants
(Myers 1985).
include those of Abrahamson et al.
(1984) and Latham (1985).
Outstanding examples of SPS include the "Big Scrub," part of the
Ocala National Forest, scrubs of the
Lake Wales Ridge, eg., the Archbold
Biological Station, and stands along
the Atlantic Coastal Ridge.
SMALL VERTEBRATE SPECIES
ASSEMBLAGES
Sand Pine Scrub
Compared with the LLP/TO sandhill
association, sand pine scrub (SPS)
has less area (250,000 ha) and a far
more limited distribution (fig. 1).
Scrub is associated with old shorelines, lake margins, and stream
courses where extremely well
washed, nutrient deficient sands
were deposited during Pleistocene
times (Kurz 1942; Laessle 1958a, b,
1967).The most widespread soils
supporting SPS are the St. Lucie,
Lakewood, and Pomello associations
(Beckenbach and Hammett 1962).
Sand pine scrub is a two-layered
community. Sand pine (Pinus clausa)
normally occurs as a relatively evenaged overstory species. The understory is comprised of 10-20 species of
evergreen shrubs 1-5m in height.
Four species of oaks comprise the
bulk of the biomass, Quercus geminata, Q. myrtifolia, Q. chapmanii, and
Q. inapina. Lesser numbers of other
species including Ceratiola ericoides,
Lyonia ferruginea, and Osmanthus
americanus add to local diversity.
Sand pine scrub is a fire climax community (Laessle 1958a, Abrahamson
et al. 1984). In contrast with LLP/TO,
SPS burns at intervals of 20-70 years;
a combination of ground and crown
fires destroys all the above-ground
vegetation. Most of the woody
plants, with the notable exception of
the sand pine and Ceratiala, readily
sprout from root crowns following
fires. Laessle (1958a), Veno (1976),
and Richardson (1977) provide data
on plant community structure of
scrubs. Recent quantitative studies
Longleaf Pinepurkey Oak
Sandhills
Amphibians and Reptiles
At least 47 species of herptiles, including 2 newts, 13 toads and frogs,
3 turtles, 10 lizards, 1 amphisbaenian, and 18 snakes, are reported
to occur in LLP/TO habitats (table 1).
Campbell and Christrnan (1982) list 5
categories of reptile and amphibian
species that occur in LLP/TO and
SPS: (1) characteristic (18 species); (2)
associated with tortoise burrows (3
species); (3) frequent (8 species); (4)
occasional (14 species); and (5) associated with aquatic habitats (21 species). Of the characteristic species, 7
are regarded as adapted to xeric conditions, 3 as sand swimmers, viz.,
Neoseps reynoldsi; Eumeces egregius,
and Tantilla relicta, and the
remainder (Sceloporus woodi, Masticophis flagellum, Stilosoma extenuaturn,
Cnemidophorus sexlineatus) to other
physical features of the habitats.
The gopher tortoise (Gopherus polyphemus) is a terrestrial turtle that
digs deep burrows in the welldrained sandhill soils (Auffenberg
and Franz 1982).Stout (1981)and
Eisenberg (1983) recognized the gopher tortoise was the keystone species in xeric pinelands. Some 80 species of animals may be classified as
burrow commensals (Cox et al. 1987);
however, the number of obligatory
commensals is much smaller. Herptiles particularly associated with gopher tortoise burrows include Rana
areolata, Pituophis melanoleucus, and
D ymarchon corais.
The snake fauna of LLP/TO
sandhills is species rich (L 18 species). This diversity includes large
forms, e.g., Dymarchon corais couperi
and Crotalus adamanteus, and small,
specialized species like Stilosoma extenuatum. This latter ophiophagous
species feeds largely on Tantilla
relicta; Tantilla, is in turn specialized
on Tenebrionidae larvae (Mushinsky
1984).
Small Mammals
At least 19 species of small mammals
with body masses less than 6.0 kg
may be anticipated in LLP/TO sandhills (table 2). Two are fossorial,
Scalopus aquatious and Geomys pinetia,
1 semi-fossorial, P. polionotus, and 2
occur in the surface litter, Blarina
carolinensis and Cyptotis parua.
Arboreal species include Sciurus
carolinensis, S. niger, Glaucomys volans,
P. gossypinus, and Ochrotomys
nuttalli. Podomys floridanus nests in
the burrows of the gopher tortoise
and the pocket gopher (Layne 1969);
it may enlarge other openings in the
soil to establish burrows independently of the gopher tortoise (R. E.
Roberts, personal observation).
Dasypus novemcinctus is the only exotic species of mammal that is clearly
established in the sandhill communi ty.
Sand Pine Scrub
Amphibians and Reptiles
Campbell and Christman (1982)
listed 64 species of reptiles and amphibians that may be found in LLP/
TO sandhills and SPS. Pitfall trapping in six different even-aged
stands of SPS on the Ocala National
Forest by Christman et al. (unpublished manuscript and personal communication) revealed 27 species
(table 1). Of 1,624 individuals
trapped, the common species were
Bufo terrestris (n=332),Cnenridophorus
sexlineatus (n=329), and Sceloporus
woodi (n=216);five species were represented by single captures. Christman et al. concluded that the herpetofaunal diversity declined with increasing age of SPS stands.
Copherus polyphemus is the keystone species in SPS but is less common there than in LLP/TO (Auffenberg and Franz 1982).Many, if not
most, of the burrow commensals are
common in SPS (Cox et al. 1987).
Podomys floridanus is an example.
Small Mammals
Fourteen species of small mammals
commonly inhabit SPS (table 2). Pod-
omys floridanus is a predictable member of the assemblage throughout the
range of scrubs in peninsular Florida
(Layne 1978).Three subspecies of
Peromyscus polionotus occur in scrubs
of the interior and east coast portions
of the peninsula. Common small
mammals in central peninsular Florida scrubs include Podomys floridanus,
Peromyscus goss ypinus, Ochrotomys
nuttalli, and Glaucomys volans (Swindell 1987).Podomys floridanus is the
predominate small mammal in
scrubs of southeast Florida
(Richardson et al. 1986).
Limited data suggest Spilogale
putorius is a major predator on small
mammals in scrubs with lesser roles
played by Mephitis mephitis and
Mustela frenata (Stout and Roberts,
personal observations).
ENDANGERED AND THREATENED
SPECIES
Ten species of amphibians, reptiles,
and small mammals associated with
xeric pineland are currently listed as
having some level of threatened, endangered, or sensitive status by either the state of Florida or the Department of Interior (table 3). The
extensive overlap in species composition between the two pineland communities results from the high number of species common to both types.
The Endangered Species Act charges
federal agencies with the responsibility to manage federally listed species
on federally owned lands. At the
state level, preservation of these
listed species is of major concern
when they occur on parcels of land
scheduled for large-scale development. Preserve design and management practices for these species have
largely evolved on an ad hoc basis
without adequate time for an evaluation of the management or the longterm implications for the species.
MANAGEMENT OF XERIC
PINELANDS ON PUBLIC LANDS
Of three national forests in Florida,
only the Ocala National Forest is located in the peninsula. It totals
153,846 ha of which 85,020 ha are SPS
and 18,219 ha LLP/TO. The National
Forest Management Act (1976) and
pursuant regulations (36 CFR 219)
require that each forest be managed
to maintain well-distributed and viable populations of wildlife species,
including species that are endangered or threatened (Norse et al.
1986).
Silvicultural systems differ between the two pineland communities. On the Ocala National Forest
sand pine scrub is routinely harvested in patchy clearcuts that range
from 16-24 ha in area. Scrub understory vegetation is allowed to regenerate naturally; however, sand pine
is seeded following site preparation
1
by a single roller chopping. The harvest rotation length is about 50 years.
In contrast, LLP/TO is ostensibly
managed on a 80-100 year rotation
and shelterwood cutting favors natural regeneration of the longleaf pine
(Don Bethancourt, personal communication). In practice, harvesting of
longleaf pine may occur in 60 years.
Effectiveness of ecosystem management in the SPS community will
be judged by the response of designated indicator species, such as gopher tortoises and scrub jays Uphelocoma coerulescens) (table 3). The gopher tortoise is also a designated indicator species for the LEP/TO community. The significance of the gopher tortoise as a keystone species
was emphasized in 1986 when harvesting of the species on national forests in Florida was made illegal
through an agreement between the
U.S. Forest Service and the Florida
Game and Fresh Water Fish Csmmission. Other species-specific management practices involving amphibians,
reptiles, or small mammals have not
been deemed necessary to carry out
on the Ocala National Forest (Don
Bethancourt, personal communication). In fact, the impact of timber
harvesting on small vertebrates of
LLP/TO and SPS communities is
simply not known.
Public lands in Florida supporting
xeric pinelands include, hut are not
limited to, state forests and state
parks. State forests with large acreages of LLP/TO, e.g., the Withlacoochee State Forest, are managed at
the ecosystem level. Prescribed burning is done every 3-8 years and future timber sales will follow a rotation length of 80-120 years; currently
rotation lengths are about 60 years
and are not regarded as favorably for
endemic wildlife. Wildlife management areas overlap the state forest
holdings and are managed for sustained yields of wildlife by the Florida Game and Fresh h7aterFish
Commission based on a memorandum of understanding between
agencies (Cathy Ryan, personal communication).
State parks are managed by the
Division of Recreation and Parks of
the Florida Department sf Natural
Resources (FDNR). An ecosystem
approach is taken in the restoration
and management of xeric pinelands
on state park lands (Jim Stevenson,
personal communication). Prescribed
burning has been used since 1969 to
control hardwood invasion of LLP/
TO stands and to stimulate growth
and flowering of grasses and herbs.
Burning in spring and early summer
appears to best duplicate the historic
timing of lightning initiated fires in
xeric pinelands. The impact of these
management practices on the plant
community has been documented
(Davis 1984); the response of reptiles,
amphibians, and small mammals is
currently under study (Stout et al.
unpublished). Generally, mature
stands of SPS have not been burned
until recently, due to the unpredictable behavior of fire in the community; however, a prescription for
burning this fuel type has been written and tested on private land and
state parks (Doran et al. 1987).Early
recovery stages of SPS appear to support the greatest diversity of reptiles
and amphibians. However, as canopy closure occurs in SPS, ground
cover diminishes and habitat quality
for gopher tortoises declines (Cox et
al. 1987). In contrast, similar numbers
of Podomys have been observed in
early (R. E. Roberts, unpublished
data, J. Dickinson State Park); intermediate (Stout 1982); and old growth
SPS (James N. Layne, unpublished
data, Archbold Biological Station).
State parks, reserves, and preserves appear to be ideal lands to explore species-specific management
measures for herptiles and small
mammals. For example, sand swimming herptiles (Smith 1982)require
openings that are relatively root free
in LLP/TO and SPS habitats. The
natural occurrence of such openings
may have been due to "hot" spots
associated with the combustion of
high fuel loads, e.g., fallen trees (Ron
Myers, personal communication).
Concentration of natural fuels prior
to prescription burns in SPS would
offer a means to create microhabitat
conditions favorable for the sand
swimmers.
MANAGEMENT OF XERIC
PINELAND ON PRIVATE LAND
Development of Regional Impact
Concern with management of amphibians, reptiles, and small mammals on private lands in Florida derives from state and federal protection of endangered species and the
development guidelines promulgated during the Development of
Regional Impact (DRI) process. "The
Florida Environmental Land and
Water Management Act of 1972"
(Chapter 380, Florida Statutes) defines developments of regional impact in Section 380.06(1),Florida Statutes, as "...any development which,
because of its character, magnitude,
or location, would have a substantial
effect upon the health, safety, or welfare of citizens of more than one
county (Anonymous 1976)." Large
scale development projects in peninsular Florida commonly involve hundreds to several thousand acres of
relatively natural landscape. The DRI
process requires bona fide studies of
wildlife populations and their associated habitats; emphasis is placed on
listed species. Developers must prepare viable management strategies to
accommodate wildlife resources dependent upon their lands (Cox et al.
1987; Richardson et al. 1986).
Management strategies of developers with xeric pinelands generally
follow one of two somewhat overlapping approaches to preserve habitat
and/or species values: (1) conservation set asides or (2) mitigation. Conservation set asides are, in principle,
the preferred solution. In practice
some habitat is dedicated in perpetuity as a nature preserve; preserve design currently is a somewhat ad hoc
process and will be discussed more
completely in a subsequent section of
this paper. Very high land values
may dictate mitigation rather than on
site preservation of habitat.
Mitigation may take many forms
to compensate for development of
xeric pinelands. Restoration of degraded land (Humphrey et al. 19851,
not necessarily xeric pinelands, is one
method. Another tactic is to purchase
comparable land or some other type
of land of equivalent natural value
elsewhere and dedicate it to preservation. A formal process for accomplishing this option is presently under study by the Florida Game and
Fresh Water Fish Commission.
Preservation of habitat is the basic
purpose of conservation set asides
and mitigations. The value of these
efforts depends on the proximity to
larger, undeveloped tracts of land,
travel corridors, area of preserves,
and future management options.
Another form of mitigation is the
relocation of sensitive species from
tracts of land to be developed to land
dedicated to purposes that are consistent with the long-term survival of
the relocated species. In Florida, the
gopher tortoise has been the focus of
numerous relocation efforts. Diemer
(1984) discussed the advantages and
disadvantages of relocation of gopher tortoises as a species management strategy. Formal research on
gopher tortoise relocation was recently reported (Proced. Gopher Tortoise Relocation Symp., 27 June 1987,
Gainesville, FL, in press). The Florida
Game and Fresh Water Fish Commission regulates relocations by a permit
system based on a standardized relocation protocol.
Preserve Design
Preserve design is an evolving and
controversial area of conservation
biology (Diamond 1975,1978; Gilbert
1980; Higgs 1981; Margules 1982;
Pickett and Thompson 1978; Pyle
1980; Soule and Simberloff 1986).
Large preserves encompassing a mo-
saic of xeric pinelands, mesic forests,
and seasonal and permanent wetlands would perhaps offer the ideal
landscape unit for long-term preservation of amphibians, reptiles, and
small mammals in peninsular Florida. Because preserves on private
lands must be justified and dedicated
through the DRI process, economics
dictates preserve units of minimal
size. Rarely do we have the opportunity to cluster or juxtapose these
small units to take advantage of the
so called "rescue effect" (Brown and
Kodric-Brown 1977).
In practice, conservation set asides
tend not only to be small in acreage
but also only of one habitat type. The
latter presents a dilemma for species
whose requirements often include
two or more contrasting habitats. For
example, the gopher frog lives in tortoise burrows in LLP/TO sandhills
during late spring, summer and early
fall and migrates to temporary wet
season depressions to breed in winter and early spring (Moler and
Franz 1987).Thus a mosaic of upland-wetland habitats in close proximity are essential to maintain viable
populations of this species. Other
species such as the indigo snake have
home range requirements that include 122-202 ha of several uplandwetland habitat types (Moler 1985;
Moler unpublished data). It is obvious that large landscape units are
necessary to preserve viable populations of these animals.
We have prepared a detailed preserve design for a SPS community
within the city of Boca Raton, Florida
(Richardson et al. 1986; Stout et al.
1987; manuscript in preparation).
The approach taken anticipated
Soule and Simberloff (1986) and recommended the area of the preserve
be sufficient to support a minimum
viable population (Franklin 1980)of
gopher tortoises because of their
status as the keystone species. Although biologically reasonable, this
basis-for determining preserve size is
often economically unrealistic from
the view point of the private land-
owner. A consortium of public landowners would, however, permit the
purchase and long-term management
of the preserve as recommended.
Cox et al. (1987) offer guidelines
for the design of preserves on private
lands to maintain gopher tortoise
populations. They employed the
computer simulation model
POPDYN (Perez-Trejo and Samson
manuscript) to determine population
viability based on different initial
sizes. Populations of 40-50 individuals were found to be likely (290%)to
persist 200 years. Based on existing
literature on home range requirements, Cox et al. (1987) recommended a minimum preserve of 10-
20 ha, depending on habitat quality,
to support 40-50 tortoises.
Another approach to determining
the area of a preserve employs "incidence functions" (Diamond 1978).
Incidence functions are species specific and derived from data sets
which reveal the fraction of plots
(discrete habitats) of different areas
that actually support the species. It is
a matter of judgement as to the
probability of occurrence, e.g. 0.5 as
opposed to 0.7, that would set a
lower limit to area for an acceptable
preserve. Data sets useful for evaluating this approach with respect to
amphibians, reptiles, and small
mammals in xeric pinelands are pres-
ently lacking. Table 4 provides data
we have gathered on area of discrete
habitats and the presence or absence
of gopher tortoises and Florida mice.
It is apparent that tortoises are less
area sensitive than Florida mice and
that Florida mice are patchy in occurrence in LLP/TO, perhaps only secondarily related to area.
Incidence functions do not necessarily reveal the minimum area required to support minimum viable
populations (Franklin 1980).We believe preserve area should be based
on providing this requirement, particularly when preserves are isolated
relative to average dispersal distances of keystone species. However,
clusters of preserves within dispersal
distances of keystone species may be
of less area per preserve due to a
high likelihood of reinvasion from
nearby populations following local
population extirpations (Noss and
Harris 1986).
Management of Preserves in Xeric
Pinelands
Cedar Grove
J. Dickinson
21.5
256.2
+
+
+
+
The future viability of preserves depends largely on their ownership after development of the surrounding
landscape. It is unlikely that homeowners associations will assume the
cost of management if preserves remain as a part of the overall development's "commons." Public ownership is an alternative and might rest
with a city, county, or state. Local
governments seem more appropriate;
however, funds and expertise to
manage may be lacking. One preserve in south Florida is designed to
border a city park, thus allowing its
maintenance and /or management
costs to be assumed over time as part
of the existing park system
(Richardson, personal observation).
Regardless of the ownership, a commi tment to long-term management
must be achieved if a preserve is to
retain natural values.
Management options for nature
preserves range from a decision I ) to
d o nothing and let nature take its
course; 2) to manage for maintenance
of a viable ecosystem, which implies
the natural biota, including amphibians, reptiles, and small mammals,
will be present in proportion to their
normal abundance; or 3) to focus
management on the needs of one or
more species. White and Bratton
(1980) have exposed the folly of the
first management option. The decision to emphasize ecosystem or species management depends on the entity responsible for management,
type of preserve, management objectives, area of the preserve, nature of
the surrounding lands, relative overall or regional rarity of particular
species, and the resources available
for management.
Management objectives of any
preserve should focus on: I) maintenance of normal ecosystem processes; 2) conservation of soil; 3)
maintenance or restoration of normal
hydrologic conditions; 4) prevention
of establishment of exotic species.; 5)
and prevention of human encroachment (e.g., dumping, ATVs, etc.) Beyond these generalities, management
of preserves is an idiosyncratic process that may concern endemic species, genetics of inbred populations,
or restoration of periodic wild fires.
Xeric pinelands of peninsular Florida depend on periodic fires to maintain their structure and function
(Laessle 1958a; Abrahamson 1984).
Thus a burning program is essential
in the management of LLP/TO or
SPS preserves. Spring or early summer prescribed burns are routinely
used to maintain LLP/TO communities on state parks. Doran et al. (1987)
have documented prescribed burns
of SPS preserves in an urban setting
based on rather esoteric fire models
developed by the U.S. Forest Service.
~ o ~ htortoises
e r
respond favorably
to the bums (Stout et al. 1988).A mosaic of recovery stages in SPS may
favor beta diversity of herptiles and
small mammals. Mushinsky (1985)
has carefully documented the response of the herpetofauna to a vari-
ety of burning schedules in LLP/TO.
Diversity and abundance of amphibians and reptiles was increased on
experimental plots relative to unburned controls. Re-establishment of
the pine overstory may be necessary
to produce needle cast for carrying
fire (Landers and Speake 1980).
Management of conservation set
a s i d s and/or easements may focus
on particular species or combinations
of species. The smaller the preserve
the more likely that a reduced suite
of species will be present
(Richardson et al. 1986).Given that a
fixed area is available for management, major efforts to enhance or
maintain habitat should target those
species that can maintain viable
gopula tions within the preserve
(Shaffer 1986). A species whose minimum area requirements for a minimum viable population exceeds the
preserve area should not be of major
concern (Shaffer and Samson 1985);
nonetheless, such species can benefit
from the preserves if travel corridors
exist (Harris 1984).
DISCUSSION
Xeric pinelands of peninsular Florida
support a species-rich assemblage of
reptiles, amphibians, and small
mammals. Growth and development
continues to diminish LLP/TO and
SPS habitats to the detriment of the
associated biota. Land in public ownership, e.g. state parks and forests,
national forests, and private holdings, e.g., the Archbold Biological
Station, and institutional lands such
as the Ordway and Swisher Preserves, jointly owned and managed
by the University of Florida and The
Nature Conservancy, will be increasingly valuable as other xeric pinelands are converted to land uses not
favorable to the biota. Thus, management of these xeric pinelands will
become more important in the future. At present management is
largely limited to prescribed bums to
maintain what were historically fire
climax communities. Thus, fire management is tantamount to small vertebrate management.
In the future as air quality standards are modified, prescribed burning, particularly in or near urbanized
areas, will be restricted or eliminated
as a management option. Alternative
means of habitat manipulation need
to be developed, particularly for SPS.
Basic information on the life history of many amphibians, reptiles,
and small mammals of xeric pinelands is lacking. The Nongarne Wildlife Program of the Florida Game and
Fresh Water Fish Commission has
initiated and f u ~ ~ d rather
ed
large
scale studies of SPS and LLP/TO
communities. These studies are at the
community level and largely observational. Management needs of individual species may be derived only
secondarily from this research. Studies that focus on particular species
will ultimately lead to more refined
habitat management guidelines. The
report by Cox et al. (1987) will likely
serve as a model for the preparation
of habitat protection guidelines; management follows protection (White
and Bratton 1980).
Management alternatives at the
ecosystem and species level are
needed now for xeric pinelands on
private lands undergoing development Regulation of development in
these habitats as currently practiced
will result in a patchwork of small,
isolated nature preserves. Preservation of natural habitat in a developed
landscape is, of course, desirable.
However, several problems remain:
(1) who will own the preserves, (2)
how will a management plan be prepared, and (3) who will be responsible for management? Even another
decade of rapid growth in peninsular
Florida may result in a feG hundred
nature preserves, which will not necessarily be restricted to xeric pineland habitat. Ignoring the question of
ownership, no public land management agency is currently capable of
assuming the charge of managing
these preserves. Lack of manage-
ment, e.g., failure to conduct prescribed burning, will allow successional changes to occur to the detriment of many small vertebrates narrowly adapted to xeric pinelands.
Loss of habitat and species values
originally used by jurisdictional
agencies to secure preserve set asides
provides a potential basis for private
land owners to request development
rights on the land. This action would
defeat the entire purpose of having
conservation set asides.
An alternative to on site habitat
protection is offered by Cox et al.
(1987) in regard to preserving habitat
for the gopher tortoise. The alternative, a Wildlife Resource Mitigation
Fund (WRMF),allows a developer to
contribute money to the fund to mitigate losses of valuable wildlife habitat on lands being developed. The
collective monies of several development projects would allow an independent group such as the Trust For
Public Lands to assist in the purchase
of commensurate lands to expand an
existing public park, preserve or forest. Management is more likely to be
applied to these lands and ultimately
the resources are better served by the
public agencies.
ACKNOWLEDGMENTS
We thank the authors of the papers
cited herein for their efforts and
dedication to science. Biologists who
contributed to our knowledge of
xeric pineland include but are not
limited to the following individuals:
Dan Austin, Don Bethancourt, Russ
Burke, Steve Christman, David Cook,
David Corey, Jim Cox, Joan Diemer,
Dick Franz, Larry Harris, Randy
Kautz, Jim Layne, Wayne Marion,
Paul Moler, Ron Myers, Reed Noss,
Cathy Ryan, Jim Stevenson, and Don
Wood. Support for research on the
ecology of sandhill communities was
provided by the Nongame Wildlife
Program, RFP86-003, of the Florida
Game and Fresh Water Fish Commission. Development groups that
funded work by the authors on xeric
pinelands include Hardy-Lieb Development Corporation, The Adler
Group, and Deutsch-Ireland Properties. The Division of Recreation and
Parks (FDNR), Department of Biology, University of South Florida, and
the Department of Biological Sciences, University of Central Florida
assisted in our studies in a variety of
ways. We thank Beverly Bamekow,
Rita Greenwell, Barbara Erwin, and
Nancy Small for typing the manuscript. Lastly, we thank Paul E.
Moler, James N. Layne and Robert C.
Szaro for providing excellent suggestions to improve the paper.
LITERATURE CITED
Abrahamson, Warren G. 1984. Postfire recovery of Florida Lake
Wales ridge vegetation. American
Journal of Botany 7l:9-21.
Abrahamson, Warren G., Ann F.
Johnson, James N. Layne, and Patricia A. Peroni. 1984. Vegetation
of the Archbold Biological Station,
Florida: an example of the Southern Lake Wales Ridge. Florida Scientist 47:209-250.
Anonymous. 1976. Developments of
regional impact: guidebook for
preparation of the application for
development approval. 65 p. Division of State Planning, Department
of Administration, Tallahassee,
Florida.
Arata, Andrew A. 1959. Effects of
burning on vegetation and rodent
populations in a longleaf pine turkey oak association in north Central Florida. Quarterly Journal
Florida Academy Sciences 22:94104.
Auffenberg, Walter, and Richard
Franz. 1982. The status and distribution of the gopher tortoise (Gopherus polyphemus). p. 95-126. In
North American tortoises: conservation and ecology. R. Bruce Bury,
editor. Wildlife Research Report
12. 126 p. U.S. Fish and Wildlife
Service, Washington, D.C.
Beckenbach, J. R., and J. W. Hammett. 1962. General soil map of
Florida. 1p. Florida Agricultural
Experiment Stations and the Soil
Conservation Service.
Brown, James H., and Astrid KodricBrown. 1977. Turnover rates in
insular biogeography: effects of
immigration on extinction. Ecology 58:445-449.
Campbell, Howard W., and Steven P.
Christman. 1982. The herpetological components of Florida sandhill
and sand pine scrub associations.
p. 163-171. In Herpetological communities: a symposium of the Society for the Study of Amphibians
and Reptiles and the Herpetologists' League, August 1977. Norman J. Scott, Jr., editor. 239 p. U.S.
Fish and Wildlife Service, Wildlife
Research Report 13.
Christrnan, Steven P., Howard I.
Kochman, Howard W. Campbell,
Charles R. Smith, and Wallace S.
Lippincott, Jr. Successional
changes in community structures:
amphibians and reptiles in Florida
sand pine scrub. Unpublished
manuscript.
Cox, James, Douglas Inkley, and
Randy Kautz, 1987. Ecology and
habitat protection needs of gopher
tortoise (Gopherus polyphemus)
populations found on lands slated
for large-scale development in
Florida. Florida Game and Fresh
Water Fish Commission Nongame
Wildlife Program Technical Report
no. 4.75 p. Tallahassee, FL.
Davis, John H. 1980. General map of
natural vegetation of Florida.
Agric. Exp. Sta. Institute of Food
and Agricultural Sciences Circular
S-178.1 p. University of Florida,
Gainesville.
Davis, Mary M. 1984. Phenology of
flowering plants on experimental
fire plots, St. Marks National
Wildlife Refuge. M. S. Thesis. 54 p.
Florida State University. Tallahassee, FL.
Diamond, Jared M. 1975. The island
dilemma: lessons of modern biogeographic studies for the design
of natural reserves. Biological
Conservation 7:129-146.
Diamond, J. M. 1978. Critical areas
for maintaining viable populations
of species. p. 2740. In The breakdown and restoration of ecosysHoldgate and M. J.
tems. M. b7.
Woodman, editors. Plenum Press,
New York, N. Y.
Diemer, Joan E. 1984. Tortoise relocation in Florida: solution or problem? Proc. 1984 Symposium Desert Tortoise Council: 131-135.
Doran, R. F., Donald R. Richardson,
and Richard E. Roberts. 1987. Prescribed burning of the sand pine
scrub community: Yamato Scrub,
a test case. Florida Scientist 50384192.
Eisenberg, J. F. 1983. The gopher tortoise as a keystone species. p. 1-4.
In The gopher tortoise: a keystone
species. R. J. Bryant and R. Franz,
editors. Proceedings of the 4th
Annual Meeting of the Gopher
Tortoise Council. Florida State
Museum, Gainesville, FL.
Franklin, Ian Robert. 1980. Evolutionary change in small populations.
p. 135-149. In Conservation biology: an evolu tionary-ecological
perspective. Michael E. Soule and
Bruce A. Wilcox, editors. 395 p.
Sinauer Assoc., Sunderland, MA.
Gilbert, F. S. 1980. The equilibrium
theory of island biogeography:
fact or fiction? Journal of Biogeography 2209-235.
Givens, Kerry T., James N. Layne,
Warren G. Abrahamson, and
Susan C. White-Schuler. 1984.
Structural changes and successional relationships of five Florida
Lake Wales Ridge plant communities. Bulletin Torrey Botanical
Club 111%-18.
Harris, Larry D. 1984. The fragmented forest. 211 p. The University of Chicago Press, Chicago,,IL.
Higgs, A. J. 1981. Island biogeography theory and nature reserve design. Journal of Biogeography
8:117-124.
Humphrey, Stephen R., John F. Eisenberg, and Richard Franz. 1985.
Possibilities for restoring wildlife
of a longleaf pine savanna in an
abandoned citrus grove. Wildlife
Society Bulletin 13:487496.
Kalisz, Paul J., and Earl L. Stone.
1984. The longleaf pine islands of
the Ocala National Forest, Florida:
a soil study. Ecology 65:1743-1754.
Kurz, Herman. 1942. Florida dunes
and scrub vegetation and geology.
Florida Geological Survey Bulletin
23:15-154.
Laessle, Albert M. 1942. The plant
communities of the Welaka Area.
University of Florida Publication,
Biological Science Series 4:l-143.
Laessle, Albert M. 1958a. The origin
and successional relationship of
sandhill vegetation and sand pine
scrub. Ecological Monographs
28:361-387.
Laessle, Albert M. 1958b. A report on
succession studies of selected
plant communities on the Univ. of
Ha. conservation reserve, Welaka,
Florida. Quarterly Journal Florida
Academy Science 21:101-112.
Laessle, A. M. 1967. Relationship of
sand pine scrub to former shore
lines. Quarterly Journal Florida
Academy Science 30:269-286.
Landers, J. Larry, and Dan W.
Speake. 1980. Management needs
of sandhill reptiles in Southern
Georgia. Annual conference
Southeastern Association Fish and
Wildlife Agencies. 34:515-529.
Latham, Pamela J. 1985. Structural
comparisons of sand pine scrubs
of east-central Florida. M. S. Thesis. 106 p. University of Central
Florida, Orlando.
Layne, James N. 1969. Nest-building
behavior in three species of deer
mice, Perornyscus. Behaviour
35288-303.
Layne, James N. 1978. Pevomyscus flon'danus (Chapman). p. 21-22. In
Rare and endangered biota of
FLorida. Vol. 1: mammals. James
N. Layne, editor. 52 p. University
Presses of Florida, Gainesville, Fl.
Margules, C., A. J. Higgs, and R. W.
Rafe. 1982. Modern biogeographic
theory: are there any lessons for
nature reserve design? Biological
Conservation 24:ll5-128.
Moler, Paul E. 1985. Distribution of
the eastern indigo snake, D ymarchon corais couperi, in Florida.
Herpetological Review 116:37-38.
Moler, Paul E., and Richard Franz.
1987. Wildlife values of small, isolated wetlands in the southeastern
coastal plain. p. 234-241. In Proc.
3rd S. E. Nongame and Endangered Wildlife Symposium. R. R.
Odom, K. A. Riddleberger and J.
C. Ozier, editors. Georgia Department Natural Resources, Game
and Fish Division.
Monk, Carl D. 1968. Successional and
environmental relationships of the
forest vegetation of North Central
Florida. The American Midland
Naturalist 79:441457.
Mushinsky, Henry R. 1984. Fire and
the Florida sand hill herpe tofaunal
community: with special attenti on
to responses of Cnemidophorus
sexlinea tus. Herpetologica 41:333342.
Myers, Ronald L. 3985. Fire and the
dynamic relationship between
Florida sandhill and sand pine
scrub vegetation. Bulletin Torrey
Botanical Club 122241-252.
Norse, Elliott A., Kenneth L. Rosenbaum, David S. Wilcove, Bruce A.
Wilcox, William H. Romme,
David W. Johnston, and Martha L.
Stout. 1986. Conserving biological
diversity in our national forests.
116 p. The Wilderness Society.
Noss, Reed F., and Larry D. Harris.
1986. Nodes, networks, and
MUMS: preserving diversity at all
scales. Environmental Management 10:299-309.
Pickett, S. T. A., and John N. Thompson. 1978. Patch dynamics
and the design of nature reserves.
Biological Conservation 13:27-37.
Pyle, Robert Michael. 1980. Management of nature reserves. p. 319327. In Conservation Biology: An
evolu tionary-ecological perspective. Michael E. Soule and Bruce
A. Wilcox, editors. 395 p. Sinauer
Associates, Sunderland, Mass.
Richardson, Donald Robert. 1977.
Vegetation of the Atlantic coastal
ridge of Palm Beach County, Florida. Florida Scientist 40:281-330.
Richardson, Donald R., I. Jack Stout,
Richard E. Roberts, Daniel F.
Austin, and Taylor R. Alexander.
1986. Design and management
recommendations for a sand pine
scrub preserve: the Yamato scrub.
124 p. Ecological Consultants,
Tampa, Florida.
Shaffer, Mark L. 1986. The metapopulation and species conservation: the special case of the northern spotted owl. p. 86-99. In Ecology and management of the spotted owl in the Pacific Northwest,
Ralph J. Gutierrez, and Andrew 8.
Carey, tech editors. USDA Forest
Service. General Technical Report
PNW-185,119 p. Pacific Northwest Forest and Range Experiment
Station, Portland, Oregon.
Shaffer, Mark L., and F. B. Samson.
1985. Population size and extinction: a note on determining critical
population sizes. American Naturalist 125:144-152.
Smith, Charles R. 1982. Food resource partitioning of fossorial
Florida reptiles. p. 173-178. In Herpetological communities: a symposium of the Society for the Study
of Amphibians and Reptiles and
the Herpetologistsf League, August 1977. Norman J. Scott, Jr.,
editor. 239 p. U.S. Fish and Wildlife Service, Wildlife Research Report 13.
Soule, Michael E., and Daniel Simberloff. 1986. What do genetics
and ecology tell us about the design of nature reserves? Biological
Conservation 35:19-40.
Stout, I. Jack. 1981. Conservation biology of Peromycus floridanus. Florida Scientist 44 (Supplement 1):37.
Stout, I. Jack. 1982. Small mammal
community structure in sand pine
scrub. Bulletin Ecological Society
America 63:68-69.
Stout, I. Jack, Donald R. Richardson,
and Richard E. Roberts. 1988. Response of resident and relocated
gopher tortoises to a prescribed
burn in a sand pine scrub community. Proced. Gopher Tortoise Relocation Symp., 27 June 1987,
Gainesville, FL.
Stout, I. Jack, Donald R. Richardson,
Richard E. Roberts, and Daniel F.
Austin. 1987. Design of a nature
preserve in a subtropical urbanizing landscape: application of ecologic and genetic principles. Bulletin Ecological Society of America
68:423-424.
Swindell, C. E. 1987. Multivariate
analysis of competition and habitat use by small mammals in Florida Sand Pine Scrub. M. S. Thesis.
59 p. University of Central Florida,
Orlando.
Veno, Patricia A. 1976. Successional
relationships of five Florida plant
communities. Ecology 57:498-508.
White, Peter S., and Susan P. Bratton.
1980. After preservation: philosophical and practical problems of
change. Biological Conservation
18:241-255.
White, W. A. 1970. The geomorphology of the Florida peninsula. Bureau of Geology, Tallahassee. 164
p. Geological Bulletin No. 51.
Wood, Don A. 1987. Official lists of
endangered and potentially endangered fauna and flora in Florida. 19 p. Florida Game and Fresh
Water Fish Commission.
Download