Title: Herpetological components of Florida sandhill and sand pine scrub associations
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Title: Herpetological components of Florida sandhill and sand pine scrub associations
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Christman, Steven P.
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The Herpetological Components of
Florida Sandhill and Sand Pine Scrub Associations


by

Howard W. Campbell' and Steven P. Christman


U.S. Fish and Wildlife Service
Denver Wildlife Research Cent,
412 N. E. 16th Avenue, Room 2
Gainesville, Florida 32601


Abstract


F~ORDR S~EMUSEU


Investigations of the herpetofauna of the peninsular Florida sandhills and sand pine
scrub revealed a diverse complex composed of a minority of xeric-adapted species com-
bined with an array of wide-ranging and aquatic species that can be found in many
Florida habitats. The xeric-adapted species required sand for burrowing or sand-swim-
ming. The tortoise (Gopherus) digs burrows that serve as shelter for several other species:
the wide-ranging generalist species, however, require none of the specialized conditions
of the sandhills and scrub. Sand scrub, especially early successional stages, contains 22
species of reptiles and amphibians, which is more than in any other of a wide range of pe-
ninsular Florida habitats. Apparently the herpetofauna is responding to the dry, well-
drained soil and patches of sand free from roots rather than to any aspect of the vegeta-
tion. The sand-swimming species (Eumeces egregius, Tantilla relicta, and Neoscps rcTy-
noldsi) depend on periodic disturbance (e. g.. fire and clear-cutting) to remove the matted
understory and pine canopy. The widespread distribution of these forms attests to the
continuous presence, throughout history, of a mix of successional stages in both the sand-
hills and sand pine scrub.


The sandhill and scrub pine plant associations
are characteristic and well defined elements of
the Florida environment. Although vegetatively
distinct, they are closely related physically and
geographically (Fig. 1) and have very similar
herpetofaunas. Autecological studies of sandhill
and scrub species are available (Telford 1959;
Jackson 1972; Smith, this volume), but, in con-
trast to extensive literature on the vegetation,
previous research has not dealt with the com-
munity of animals living in these plant associa-
tions. The sandhills and sand pine scrubs, along
with xeric hammocks, form the principal habi-
tats in Florida for a distinctive herpetofauna
based on xeric-adapted forms.
We initiated studies (Florida Fish and Game
Commission 1976) in these habitats in the spring
of 1975 using the methods detailed elsewhere
(Campbell and Christman, this volume). In 1976

IDeceased: Died at Gainesville, Florida. on 10 Decem-
ber 1981.


we substantially increased our efforts in the
various subtypes of sandhill and scrub in the
Ocala National Forest in central Florida to deter-
mine the distribution and relative abundance of
amphibian and reptile species in the various suc-
cessional stages of sandhills and scrub.


Characterization of the Habitats

Sandhill Association

The sandhill association and its successional
relations to other Florida plant associations have
received considerable attention over the years
(Laessle 1958), and there is general agreement
on the physical and phytogeographic character-
istics of the sandhills. References to the reptiles
and amphibians occurring in the sandhills gen-
erally use the designators "High Pine" (Carr
1940) or "Longleaf-pine/Turkey-oak" (Laessle
1942). Similar habitats, differing only slightly in






H. W. CAMPBELL AND S. P. CHRISTMAN


.' -i..


Fig. 1. Distribution of sandhills (hatched) and sand pine scrub (black) in Florida (Davis 1967).


their plant species, are found in scattered areas
having suitable soils throughout the southern
United States (Laessle 1958; Bozeman 1971),
but the sandhills of the Florida peninsula sup-
port a herpetological community that is distinct
from the sandhills of the Florida panhandle and
other areas of the southeast. The following
analysis will deal specifically with the peninsu-
lar sandhills.
Most sandhills in peninsular Florida occur on
well-drained, sandy soils of the Lakeland series
as defined by Gammon et al. (1953) and Laessle
(1958). The physical characteristics of these soils
and their similarity to the soils supporting sand
pine scrub and xeric hammock, as well as the


relative frequency of fire in these associations,
are considered to be the factors determining
which animals use these habitats. Sandhills are
generally restricted to soils of slightly higher clay
and silt content than those supporting the scrub
association (Laessle 1958).
The sandhill association exists in two basic
forms in peninsular Florida: the "typical" long-
leaf pine (Pinus palustris)/turkey oak (Quercus
laevis) association, and the turkey oak sandhills.
The typical phase is a three-tiered habitat with
widely-spaced longleaf pines forming the upper
level, an understory of scattered turkey oak
(bluejack oak, Q. incana, may replace turkey
oak on lower soils of the Blanton series), and a





HERPETOFAUNA OF FLORIDA SANDHILLS AND SCRUB


Fig. 2. Typical phase of the longleaf pine-turkey oak sandhill habitat in the Ocala National Forest, Florida.


ground cover dominated by wire grass (Aristida
stricta and Sporobolus gracilis. Fig. 2). Fire is
frequent in this habitat, perhaps the most wide-
spread of the fire subclimax associations in
Florida. The dominant plant species are fire re-
sistant.
Where the longleaf pines are removed, the
turkey oak increases in height, forming the can-
opy layer at 8-10 m, and the ground cover be-
comes less dense with frequent and extensive
areas of bare sand interspersed with drifted piles
of oak leaves and scattered vegetation (Fig. 3).
Fire is also a frequent phenomenon in the turkey
oak sandhill and often results in extensive areas
of open sand and occasional damage to the oaks.
The turkey oak sandhill is by far the predomi-
nant phase of sandhill habitat remaining in pe-
ninsular Florida; lumbering demands on the
longleaf pine long ago eliminated it over ex-
tensive areas of sandhill habitat. Today the
Riverside Island area of the Ocala National For-
est (Fig. 2) is perhaps the largest and least dis-
turbed stand of the "typical" longleaf pine-dom-
inated sandhill habitat remaining in the south-
eastern United States (Florida Game and Fresh
Water Fish Commission 1976).


Sand Pine Scrub Association

The sand pine scrub association occurs
throughout Florida in isolated patches, often as-
sociated with or adjacent to sandhills (Fig. 1).
The soils that support scrub habitat are of the St.
Lucie series and are essentially loose, unconsoli-
dated sands, similar to but coarser and looser
than those which support sandhills. There is
never standing or running water in scrub habi-
tats, as even the heaviest rains percolate imme-
diately into the loose sand. Mulvania (1931:
528) referred to these soils as ". .. a bed of sil-
ica, to which the term soil is but remotely appli-
cable." The Florida scrub has been called the
ecological equivalent of the California chapar-
ral (Laessle 1967).
Botanists have been intrigued with scrub since
its description early in the last century (Vignoles
1823). Perhaps no plant community in Florida
has stimulated more interest or more printed
words (e.g., Vignoles 1823; Nash 1895: Whitney
1898; Harper 1914. 1921; Mulvania 1931; Web-
ber 1935; Kurz 1942; Laessle 1942, 1958, 1967;
Miller 1950; Cooper et al. 1959; Veno 1976).
Characteristic scrub tree species include the






H. W. CAMPBELL AND S. P. CHRISTMAN


Fig. 3. Turkey oak phase of tlhe longlaf pine-turkey oak sandhill habitat in the Ocala National Forest. Florida.


sand pine (Pinuts clausa), various evergreen oaks
(Quercus virginiana. Q. myrtifolia, and Q.
chapmani, and lyonia (Lyonia ferrugtinca).
Scrubs can be divided into two major types:
those with sand pine trees and those without.
The latter are sometimes referred to as rosemary
scrubs. It is not always clear why some scrubs
lack sand pines. Where the pines have died of
old age in the absence of fire, scrubs sometimes
persist for many years (Laessle 1967). Other
evergreen oak scrubs appear never to have had
sand pines. The Florida scrub jay (Apheloroma
coerule.scens), the sand skink (Neosep7s rey-
noldsi), and the Florida scrub lizard (Sceloporus
troodi) are largely restricted to the evergreen
oak scrub without sand pines and to those scrubs
with young sand pines.
Scrubs rarely burn but, when they do, the fire
usually crowns, killing the sand pines. Under the
extreme heat of a crowning fire, the sand pine
seeds are released from the serotinous cones.
More often than not, scrubs are adjacent to
sandhill communities which burn frequently.


Many authors have noted the tendency of
ground fire in the sandhills to stop abruptly at
the sandhill scrub ecotone (e.g., Webber 1935;
Laessle 1967). The dominant sandhill tree,
longleaf pine, is well adapted to survive
frequent ground fires which are readily kindled
by the wire grass and deciduous oak leaves
which accumulate rapidly. On the other hand.
the evergreen oaks of the scrub provide little fuel
for ground fires. Thus Webber (1935) referred to
the scrub as a "fire fighting association." If fire is
excluded from the scrub, the sand pines
eventually die of old age and the habitat may
ultimately succeed to hardwood (oak, magnolia)
hammock (Veno 1976). When a scrub burns.
however, the sand pines are killed, seeds are
released, and the cycle begins again. At first
there are extensive areas of open sand, but the
vegetation gradually fills in to form an almost
impenetrable tangle composed of a matted
ground cover, a dense evergreen shrub layer,
and a full pine canopy characteristic of a mature
scrub.


i4 +'':
c='ryr ;:;.~
'E' .-






HERPETOFAUNA OF FLORIDA SANDHILLS AND SCRUB


Table 1. Categories of reptile and amphibian species occurring in sandhills and scrubs in
peninsular Florida.


Characteristic
Xeric adapted
Eumeces egregious
Neos7 s reinoldsi
Sceloporius woodi
Masticophis flagellum
Stilosoma extenuatum
Tantilla relicta
Cnemidophoruts sexlineatus
Wide-ranging
Scaphiopus holbrooki
Gastrophryne carolinensis
Bufo quercicus
B. terrestris
Rhineura floridana
Anolis carolinensLi
Eumeces inexpectatus
Scincella lateral
Cemphora cocrinea
Coluber constrictor
Micrurus fulvimrs


Associated with
Tortoise Burrows
Rana areolata
Gophers polyphemus
Pituophis melanoleucms


Frequent
Hyla femoralis
It. gratiosa
Ophisaurus compresses
Diadophis punctatus
Heterodon platyrhinos
Lampropeltis triangulumn
Opheodrys aestivus
Sistruris miliarious



Results

General

Several distinct groups of amphibian and rep-
tile species occur in the sandhills and scrub
(Table 1): those that can be considered to be
highly adapted to a xeric, sandy habitat (i.e.,
reaching maximum population levels or found
only there); those that occur throughout a wide
habitat spectrum; those species associated with
burrows of tortoise (Gopherus polyphcmus);
and those species that occur in or near aquatic


Occasional
Hyla cinerea
H. crucifer
H. squirella
Pseudacris ornata
Ophisaurus attenuatus
0. ventralis
Sceloporus undulatu4s
Drymarchon corais
Elaphe guttata
E. obsoleta
Heterodon simnus
Storeria occipitomaculata
Thamnophis sirtalis
Crotalus adamanteus
Associated with
Aquatic Habitats
Acris gryllus
Rana catesbeiana
R. grylio
R. sphenorcphala
Ambystoma talpoideum .
A. tigrinumn
Notophthalmus perstriattus
N. viridescens
Pseudobranchus striatus
Siren intermedia
S. lacertina
( I,r, ,,,< floridana
C. nelsoni
Deirochelys reticularia
Kinosternon bauri
K. subrrubrum
Sternotherus odoratus
Nerodia fasciata
Seminatrix pygaea
Thamnophis sauritus
Agkistrodon pisc(iorus



habitats surrounded by sandhills, scrub, or any
other terrestrial habitat.
All of the xeric-adapted species except the
Florida scrub lizard require loose, well-drained
sand for burrowing or sand-swimming. Go-
pherus and associated species are also dependent
on sandy soils for burrows. These species are re-
stricted to sandhill, scrub, and xeric hammock
habitats, although Tantilla relicta may rarely
occur in more mesic habitats. Gopherus poly-
phemus deserves special mention as a focal spe-
cies in providing burrow refuges for a wide va-
riety of other species including Rana areolata






H. W. CAMPBELL AND S. P. CHRISTMAN


and Pituophis melanoleutLs, which are essen-
tially restricted to this microhabitat in peninsu-
lar Florida. Drymarchon corais, generally con-
sidered a characteristic gopher tortoise burrow
inhabitant, is not so restricted in peninsular
Florida and may actually reach greater popula-
tion levels in certain more mesic hardwood hab-
itats. Its close relation to gopher tortoise bur-
rows and xeric habitats is encountered primarily
at its range margins in Georgia and Alabama, in
peninsular Florida the burrows are more com-
monly used by Pituophis and a variety of other
snake species.
The aquatic species occurring in the sandhills
and scrub may reach high population levels
around appropriate habitat but are best con-
sidered fortuitous additions to the species list.
The greatest number of species occurring in the
sandhill and scrub habitats are ecological gener-
alists which find suitable conditions in a wide
variety of habitats. Unfortunately, we know too
little of absolute population densities of amphib-
ian and reptile species to be able to judge rela-
tive habitat quality for most species and thus
cannot rank the value of sandhills and scrub to
these generalist species. On inspection of Table
1, however, it is clear that the generalists are all
species which require none of the specialized
physical or vegetative conditions of the sandhill
and scrub associations.



Scrub Species Diversity

The community of herpetologists in Florida
has long been familiar with the scrub endemics
(Neoseps reynoldsi and Sceloporous woodi), but
when our data showed that scrubs had the high-
est amphibian and reptile species richness of 11
terrestrial habitat types sampled (Table 2), we
decided to look more closely at what Carr
(1940:8) called, "... undoubtedly the most rig-
orous habitat in Florida .
We list 43 nonaquatic species of amphibians
and reptiles as known to occur either character-
istically, frequently, or occasionally in Florida
scrubs and sandhills (Table 1). Two of these,
N'eoseps renoldlsi and Sceloporus woodi. are
primarily restricted to scrub habitat. Others
(e.g., Eunieces egregius, Eiumeces inexpectatis,.
Cemophora coccinea. Tantilla relicta) may
reach their greatest abundance in some succes-
sional stage of the scrub habitat. Still others oc-


Table 2. Results of the herpetological trapping
program by habitat type, Cross Florida Barge
Canal Wildlife Restudy. Eleutherodactylus
planirostris, an exotic species, has been
omitted.


Total
number
Habitat type of species


River swamp
Slash pine fatw\oods
Longleaf pine
flatwoods
Pond pine flatwoods
IHydric hammock
MNesic hanunock
Xeric lammunock
I.ongleaf pine sandhills
Deciduous oak
sandhills
Evergreen oak scrub
Sand pine scrub
(mature)


3
7


Number of
individuals/arrav/
day
0.05
0.07

0.13
0.17
0.23
0.21
0.15
0.22


16 0.11
21 0.16

19 0.14


cur in scrub only near its ecotone with sandhills
or along roads through the scrub (e.g., Go-
pherus polyphemrus and its symbionts).
The reptiles that inhabit the scrub proper fall
into two categories: those that "swim" beneath
the sand surface (Mosauer 1932) and those that
run on the surface of the sand. Even lizards such
as Anolis carolineisis and Sceloporus troodi
spend much time on the surface, as pit-fall rec-
ords indicate. Neoseps reynoldsi, Eumeces egre-
ginds, and Tantilla relicta are the principal sand
swimmers, although other small species are fre-
quently encountered beneath the sand surface
(e.g., Scincella lateral, Lammpropeltis triangu-
lui, Illeterodon platyrhinos). Smith (this vol-
ume) investigated the details of the niches of the
three principal sand-swimming species.
Amphibians are poorly represented in the
scrub. Notophthlalmnus perstriatus efts occasion-
ally inhabit the scrub. Only the most xeric-
adapted anurans will be found regularly in true
scrub. Turtles are essentially lacking, although
Kinosternon bauri may occasionally wander
through scrubs, and Copherus polyphemus will
occur near roads and ecotones where the soil
will sustain burrow construction. Snake and liz-
ard diversity is high.






HERPETOFAUNA OF FLORIDA SANDHILLS AND SCRUB


El
iL
Lol



I-
_I
0r


Fig. 4. Total numbers of species and individuals of
reptiles and amphibians collected in various ages of
sand pine scrub in the Ocala National Forest, Flor-
ida.


Scrub and Sandhill Succession

Our ongoing trapping program in the Ocala
National Forest was designed to determine if
there are any differences in the terrestrial herpe-
tofauna between different-aged stands of sand
pine scrub and between the facies of the sandhill
association. We installed two standardized trap-
ping arrays in turkey oak and longleaf pine
sandhills and in each of six various-aged stands
of even-aged sand pine. During the first 10
months of this sampling, we collected 891 speci-
mens of 22 species in the scrub traps (Fig. 4).
These results suggest strongly that some species
are more abundant in the younger stands of
scrub while others are more abundant in the
more mature scrubs. No species is confined to
the more mature scrubs, although several species
collected in the younger stands have not been
taken in the mature scrub. Of 22 species col-
lected, only Anolis carolinensis, Scincella later-
ale, and possibly Eumeces inexpectatus are more
abundant in the older scrubs. On the other
hand, at least six species occur more frequently
in the younger scrubs. Notable among these are
Scaphiopus holbrooki, Cnemidophorus sexlinea-
tus, Sceloporus woodi, and Tantilla relicta. The
1-year, 2-year and 4-year-old sites have thus far
produced 593 specimens of 20 species, whereas
the 6-year, 15-year, and mature sites have
yielded only 298 specimens of 16 species. A simi-
lar pattern characterized the turkey oak and
longleaf pine sandhills. We believe that addi-


SPECIES


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a

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,Ci


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tional collecting will not appreciably change
this trend; that is, it appears that species richness
and animal abundance decrease as the pines ma-
ture or, to be more specific, as the ground cover
increases.



Discussion

It appears to us that the amphibian and rep-
tile fauna occurring in the sandhills and scrub is
not actually responding to a particular plant as-
sociation, but rather to the physical characteris-
tics of the habitat. Where these physical charac-
teristics are met in other plant associations (e.g.,
xeric hammock), many of the same vertebrate
species occur. Thus the scrub lizard and the sand
skink are really adapted to habitats that are dry,
well-drained, and offer patches of open sand,
free from rooted vegetation. We can think of no
amphibian or reptile species in Florida that has
a distribution restricted to a single plant associa-
tion. Actually, we argue that the plant species
occurring in a given habitat are responding to
many of the same environmental factors as the
animals. Perhaps wildlife habitats should be
classified according to the significant physical
characteristics to which both the plants and the
animals are adapted rather than by plant asso-
ciations. The present system can lead to mislead-
ing judgments, and we offer a single example:
the Florida gopher tortoise is known from sev-
eral "habitats" (i.e., plant associations): longleaf
pine sandhills, deciduous oak sandhills, ever-
green oak scrub, sand pine scrub, xeric ham-
mock, dry flatwoods, and a host of ruderal sit-
uations. Does this mean that the gopher tortoise
is a highly-adaptable, ubiquitous species? It does
not. Gophers are essentially restricted to habi-
tats with well-drained, sandy soils, and an
abundance of grasses and forbs, and ultimately,
with maximal light intensity at ground level
(Auffenberg and Franz 1980). These conditions
can be met in a variety of places regardless of the
specific plant species living there.
A comparison between longleaf pine and tur-
key oak sandhill may shed light on the evolution
of the highly adapted sandhill and scrub reptile
and amphibian species. Aside from the presence
or absence of a pine canopy and the correspond-
ing dwarfing or enlargement of the understory
oaks, these facies of the association are strikingly
different in the nature of the ground cover. The


S- A . k . ktR






H. W. CAMPBELL AND S. P. CHRISTMAN


longleaf pine-dominated sandhill usually sup-
ports a dense ground cover of wire grass and
forbs, whereas the turkey oak-dominated facies
is more open, often with extensive areas of bare
sand, and it receives more intensive surface in-
solation. The reptiles characteristic of the sand-
hills are, as noted, species which burrow and
highly specialized sand "swimmers." Species
such as Tantilla relicta, Eumeces egregious. and
Neoseps reynoldsi. which require loose sand for
sand swimming, historically have encountered
such conditions in the sandhills only in areas of
disturbance, such as the burrow mounds of Go-
pherts polyphemus and Geomys pinetis. or the
open areas resulting from fire, especially under a
turkey oak canopy. In the longleaf pine/turkey
oak association such conditions (minus man's
imposition of road shoulders and other disturb-
ances) are localized islands of loose sand pro-
truding from a cover of wire grass. Similarly,
the mat of roots and humus covering the sand in
the mature sand pine scrub would require
breaks to provide suitable conditions for these
species.
Neoseps reynoldsi, Eumeces egregious. Tan-
tilla relicta, and Stilosoma extenuatum are also
more abundant in early successional stages of
sand pine scrub than they are in the advanced
stages with a full pine canopy, dense evergreen
shrub layer, and matted ground cover. To pro-
vide conditions for the evolution of the endemic
or characteristic species of the sandhills and
scrub, there must have always been such dis-
turbed areas where loose surface sand was avail-
able. Despite the contention that the central
sandhills of Florida were historically blanketed
by a sheet of longleaf pine/turkey oak (Laessle
1958), it would appear more probable that a
mix of successional stages, both in sandhills and
sand pine scrub, has always existed. Breaks in
the typical or mature stages of both habitats
could, and probably did, result from severe
fires, tornado or hurricane blow-downs of the
canopy trees, outbreaks of pine-bark beetles, or
other natural disasters. These factors would
have maintained a mix of habitats amply sup-
plied with the open, loose sand conditions
needed for the evolution of the unique herpeto-
logical components of the peninsular Florida
sandhill and scrub associations. These species,
especially Neoseps reynoldsi, Tantilla relicta,
and to a lesser extent, Eumeces egregius, are
thus cast as "weed" species, colonizers of a


patchy early successional or disturbed habitat
type which occurs throughout the sandhill, sand
pine scrub, and xeric hammock vegetative asso-
ciations as a result of biological (Gopherus. etc.)
or catastrophic (fire) factors.
Although most of the highly adapted scrub
and sandhill fauna meet few or none of the bio-
logical requirements typically cited for coloniz-
ing species (see Baker and Stebbins 1965), they
must be considered colonizers because of the
short-lived nature of their required habitat. A
population of sand skinks cannot persist to its
maturity in a patch of sand pine scrub; they in-
vade young scrubs opened by killing fires, scrubs
without sand pine, and sandhills without wire
grass and longleaf pines. Just how a species with
a reproductive rate of only one or two young per
female per year (Telford 1959) and such ap-
parently poor vagility can do this is a question
worth pursuing.
In our current study of the scrubs of the Ocala
National Forest, we consistently trapped more
species and more individuals in the younger
sand pine scrubs. This suggests that there are
either more animals in the younger stands or
that those present are easier to trap. It is clear
that the recently clear-cut scrub is more fre-
quently used by "typical" scrub species; when
we consider the natural history of scrub (i.e.,
fire ecology), this makes sense. It should come as
no surprise that the scrub-adapted fauna is ac-
customed to survival in even-aged stands of sand
pine. It thus appears that clear-cutting and
even-age management of sand pine stands may
mimic the natural situation of infrequent crown
fire to which the scrub fauna is adapted.
The variables of a herpetological association
or "community" in the scrub and sandhills habi-
tat are difficult to define. Many of the most
common species are broadly distributed and
only fortuitously juxtaposed in the sandhills and
scrub; the more specialized species occur only in
restricted microhabitats. With only a few excep-
tions, the species list for these areas can be dupli-
cated in various other habitat types in peninsu-
lar Florida. If we exclude aquatic habitats, the
species list is virtually identical to that for xeric
hammock.
It appears evident from these data that the
concept of a herpetological association or "com-
munity" cannot be defined along plant associa-
tion boundaries in the xeric habitats of peninsu-
lar Florida. The local distribution of species is






HERPETOFAUNA OF FLORIDA SANDHILLS AND SCRUB


determined by a complex of physical and proba-
bility factors at least partially independent of
the plant species of the area. The most heuristic
approach is to view amphibian, reptile, and
plant species as responding to a similar set of
physical and biotic factors without assuming
any interdependency between species. If the def-
inition of a herpetological "community" is to be
more than the fortuitous association of a group
of individuals of a variety of species at a specific
place and time, we must look beyond the plant
association. In the xeric habitats of peninsular
Florida, the particular plant associations appear
less important than the presence or absence of
areas of loose dry sand.


Acknowledgments

The staff of the Ocala National Forest have
been of immeasurable assistance to our studies at
all levels, from the initial planning stages to the
frequent extrication of vehicles from beds of soft
sand. Mrs. W. Guerrero, B. Knowles, and R.
Roth should especially be mentioned. C. R.
Smith deserves special thanks for his faithful
efforts in running the trap lines under the best
and worst of conditions. Howard Kochman and
W. S. Lippincott helped in a variety of
important capacities throughout the study.


References

Auffenberg, W., and R. Franz. 1981. The status and
distribution of Gopherus polyphemus. In R. B. Bury,
ed. North American tortoises: conservation and
ecology. In press.
Baker, H. G., and G. L. Stebbins [eds.]. 1965. The ge-
netics of colonizing species. Academic Press. New
York. 588 pp.
Bozeman, J. R. 1971. A sociologic and geographic
study of the sand ridge vegetation in the Coastal
Plain of Georgia. Ph.D. Thesis, University of
North Carolina, Chapel Hill. 243 pp.
Carr, A. F. 1940. A contribution to the herpetology
of Florida. Biol. Sci. Ser.. University of
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