• TABLE OF CONTENTS
HIDE
 Front Cover
 Synopsis
 Introduction
 Acknowledgement
 Area of study
 Physiography and geology
 Climate
 Vegetation and animal habitats
 Composition of the herpetofaun...
 Ecology of the herpetofauna
 Geography of the herpetofauna
 Origin of the herpetofauna
 Accounts of the species
 Summary
 Literature cited
 Back Cover






Group Title: Amphibians and reptiles of southern Florida (FLMNH Bulletin v.3, no.5)
Title: Amphibians and reptiles of southern Florida (FSM Bulletin, vol. 3., no. 5)
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Permanent Link: http://ufdc.ufl.edu/UF00000978/00001
 Material Information
Title: Amphibians and reptiles of southern Florida (FSM Bulletin, vol. 3., no. 5)
Series Title: Amphibians and reptiles of southern Florida (FSM Bulletin)
Physical Description: Book
Creator: Duellman, William Edward
Schwartz, Albert
Publisher: Bulletin of the Florida State Museum of Biological Sciences, University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1958
 Record Information
Bibliographic ID: UF00000978
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: ltqf - AAA0354
ltuf - ACK0923
alephbibnum - 000440457
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Table of Contents
    Front Cover
        Front Cover 1
        Front Cover 2
    Synopsis
        Page 181
    Introduction
        Page 182
    Acknowledgement
        Page 183
    Area of study
        Page 184
    Physiography and geology
        Page 184
        Page 185
        Page 186
        Page 187
    Climate
        Page 188
    Vegetation and animal habitats
        Page 188
        Page 189
        Page 190
        Page 191
        Page 192
        Page 193
        Page 194
        Page 195
        Page 196
        Page 197
        Page 198
        Page 199
        Page 200
        Page 201
        Page 202
    Composition of the herpetofauna
        Page 203
        Page 204
        Page 205
    Ecology of the herpetofauna
        Page 206
        Page 207
        Page 208
        Page 209
        Page 210
        Page 211
        Page 212
    Geography of the herpetofauna
        Page 213
        Page 214
    Origin of the herpetofauna
        Page 215
        Page 216
        Page 217
        Page 218
        Page 219
        Page 220
        Page 221
    Accounts of the species
        Page 222
        Page 223
        Page 224
        Page 225
        Page 226
        Page 227
        Page 228
        Page 229
        Page 230
        Page 231
        Page 232
        Page 233
        Page 234
        Page 235
        Page 236
        Page 237
        Page 238
        Page 239
        Page 240
        Page 241
        Page 242
        Page 243
        Page 244
        Page 245
        Page 246
        Page 247
        Page 248
        Page 249
        Page 250
        Page 251
        Page 252
        Page 253
        Page 254
        Page 255
        Page 256
        Page 257
        Page 258
        Page 259
        Page 260
        Page 261
        Page 262
        Page 263
        Page 264
        Page 265
        Page 266
        Page 267
        Page 268
        Page 269
        Page 270
        Page 271
        Page 272
        Page 273
        Page 274
        Page 275
        Page 276
        Page 277
        Page 278
        Page 279
        Page 280
        Page 281
        Page 282
        Page 283
        Page 284
        Page 285
        Page 286
        Page 287
        Page 288
        Page 289
        Page 290
        Page 291
        Page 292
        Page 293
        Page 294
        Page 295
        Page 296
        Page 297
        Page 298
        Page 299
        Page 300
        Page 301
        Page 302
        Page 303
        Page 304
        Page 305
        Page 306
        Page 307
        Page 308
        Page 309
        Page 310
        Page 311
        Page 312
        Page 313
        Page 314
        Page 315
        Page 316
        Page 317
    Summary
        Page 318
    Literature cited
        Page 319
        Page 320
        Page 321
        Page 322
        Page 323
        Page 324
    Back Cover
        Page 325
        Page 326
Full Text
mi -


BULLETIN

OF THE


FLORIDA STATE MUSEUM


BIOLOGICAL SCIENCES


Volume 3


Number 5


AMPHIBIANS AND REPTILES OF SOUTHERN FLORIDA


William E. Duellman and Albert Schwartz











UNIVERSITY OF FLORIDA
Gainesville
1958


= W, I










The numbers of THE BULLETIN OF THE FLORIDA STATE MUSEUM,
BIOLOGICAL SCIENCES, will be published at irregular intervals. Volumes
will contain about 300 pages and will not necessarily be completed in any one
calendar year.




















WILLIAM J. RIEMER, Editor
ROLAND F. HUSSEY, Associate Editor




















All communications concerning purchase or exchange of the publication should
be addressed to the Curator of Biological Sciences, Florida State Museum, Seagle
Building, Gainesville, Florida. Manuscripts should be sent to the Editor of the
BULLETIN, Flint Hall, University of Florida, Gainesville, Florida.


Published 10 December 1958


II


Price for this issue $1.63














AMPHIBIANS AND REPTILES

i OF SOUTHERN FLORIDA



WILLIAM E. DUELLMAN

and

ALBERT SCHWARTZ 1




SYNOPSIS: The 85 species of reptiles and amphibians known to occur in
southern peninsular Florida and the Florida Keys are treated systematically. The
discussion for each species includes data on variation, ecology, life history, habits,
and distribution. The definition of geographic races in southern Florida has
been accomplished only after extensive analyses of the variation encountered
throughout the peninsula. The area of study is broken down into six major habi-
tats; the distribution of the species in these habitats is analyzed. Ideas on the
origin and relationships of the herpetofauna of southern Florida are assayed on
the basis of a knowledge of the history of the peninsula and Florida Keys and on
what is known of the relationships of the component species of the herpetofauna.
TABLE OF CONTENTS
Introduction 182
Acknowledgments .183
Area of study 184
Physiography and geology.. . 184
Climate 188
Vegetation and animal habitats 188
Composition of the herpetofauna 203
Ecology of the herpetofauna 206
Geography of the herpetofauna .. . 213
Origin of the herpetofauna .215
Accounts of the species..... . 222
S Summary. . 318
Literature cited . .. .. 319

1William E. Duellman is an Instructor in the Department of Biology at
Wayne State University, Detroit, Michigan. (This paper is contribution No. 13
from that department.) He also is associated with the Museum of Zoology at the
University of Michigan, where the work on this study was done; he completed
the study under a grant from the National Academy of Sciences in 1957. Albert
Schwartz is an Instructor in the Department of Biology at Albright College,
S Reading, Pennsylvania. The collections for the present study were made while
he was a student at the Museum of Zoology at the University of Michigan. Man-
uscript submitted 14 January 1958.-ED.


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BULLETIN FLORIDA STATE MUSEUM


Vol. 3


INTRODUCTION
The southern tip of the Florida peninsula with its adjoining chain
of islands, the Florida Keys, represents the only area in continental
United States wherein semitropical and even tropical conditions pre-
vail. The peninsula also represents a cul-de-sac for animals from
the north and a springboard for those from the West Indies. How-
ever, man has also moved into southern Florida in great numbers,
and in recent years he has wrought destruction to many of the natural
habitats, so that many square miles of the "Big Cypress" have passed
through the sawmill and no longer provide a mysterious abode for
the "lurking denizens of the swamp." The Everglades have been
burned and drained, and once where great bull alligators bellowed
at night and huge flocks of snowy egrets and roseate spoonbills fed
in the marshes by day, large herds of Brahman cattle now dot the
landscape. In places oil derricks interrupt the flatness of the saw-
grass prairies and stand as monuments to human exploitation. Many
of the most beautiful and extensive tropical hammocks in the Miami
area have been hewed away to make room for homes. Already it
is too late to learn of many of the intimate relations between the
animals and their natural environment.
Our original intention was to present the accumulated data on
the amphibians and reptiles of southern Florida in a manner that
would be usable not only for the herpetologist, but for the layman
as well. With the appearance of Carr and Goin's "Guide to the rep-
tiles, amphibians, and fresh-water fishes of Florida" in 1955, this plan
was abandoned; consequently, the material is embodied in this techni-
cal report.
This study grew out of the collections of amphibians and reptiles
made by the junior author while he was studying the mammals of
southern Florida. He spent the springs and summers of 1951 and
1952 in that area. Even a cursory examination of his extensive col-
lections showed the need for additional material and extensive field
work to untangle many problems concerning the herpetofauna. Both
of us worked in the field in the summer of 1953, during which time
we attempted to collect adequate series of many species and to de-
termine the geographical distribution of all of the forms. Our field
work was repeated in the summer of 1954, at which time the em-
phasis was placed on gathering ecological data. The only other
major field work was a winter excursion to the Florida Keys by the
junior author and L. Neil Bell in 1953. The field work resulted in
the collection of several thousand herpetological specimens, of which


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 183

4740 were deposited in the University of Michigan Museum of
Zoology; smaller numbers were deposited in the University of
Miami reference collection and in the Charleston Museum. These
added to previously existing specimens in various museums in the
United States make a total of 8562 specimens from southern Florida
that were examined during the course of this study.

ACKNOWLEDGMENTS
While the authors were in the field they enlisted the aid of several
people for collecting specimens and called on others for information
and observations. Although the entire list is too long to print here,
each individual has our sincere thanks for the time and effort that
helped to make this study a reality. Certain individuals must be
singled out for special acknowledgment. Among these are three
residents of Miami, Florida-L. Neil Bell, Dennis R. Paulson, and
Raymond P. Porter. Bell, who at one time was working with the
authors on this report, was responsible for the collection of large
numbers of specimens and for gathering data on the distribution and
life histories of many species. Paulson, through his energetic field
work, greatly enlarged our collections. Porter, with his knowledge
of Florida wildlife and the Everglades, was a constant source of
valuable information and ideas. To these people we are deeply in-
debted, for had it not been for their interest and enthusiasm, much
of the data presented here would still be awaiting our discovery.
Other individuals deserving special mention for their assistance in
the field are Ann S. Duellman, Julian Harrison, Norman Hartweg,
James E. Mosimann, Daniel R. Stanland, Thomas M. Uzzell, Jr.,
and Harold J. Walter.
For the loan of specimens or permission to work in their respec-
tive institutions we wish to thank Charles M. Bogert, American Mu-
scum of Natural History; Doris M. Cochran, United States National
Museum; the late Emmett R. Dunn, Acadamy of Natural Sciences of
Philadelphia; Coleman J. Goin and Duke Wilder, University of Flor-
ida; Arthur Loveridge, Museum of Comparative Zoology; Wilfred
T. Neill, Ross Allen Reptile Institute; M. Graham Netting and Neil
Richmond, Carnegie Museum; Oscar T. Owre, University of Miami;
William J. Hamilton, Jr. and C. Richard Robins, Cornell University;
the late Karl P. Schmidt, Chicago Natural History Museum; and
Hobart M. Smith, University of Illinois. In one way or another
Walter Auffenberg, Roger Conant, Richard Edgren, Arnold B. Grob-
man, William Haast, and M. B. Mittleman have helped with certain


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BULLETIN FLORIDA STATE MUSEUM


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problems. Irving J. Cantrall, Fred G. Thompson, and Harold J.
Walters of the University of Michigan Museum of Zoology, and
H. F. Strohecker of the University of Miami, identified invertebrate
remains from stomach contents. Pierre Dansereau, formerly of the
University of Michigan and now of the Universite de Montreal, greatly
assisted the senior author in formulating the ecological section of this
report; a preliminary paper on this aspect of the study was presented
at the meetings of the American Institute of Biological Sciences and
the American Society of Ichthyologists and Herpetologists in Gaines-
ville, Florida, in 1954.
All of the senior author's work on this project was done at the
University of Michigan Museum of Zoology. Sincere thanks are due
to Norman Hartweg, T. H. Hubbell, the late J. Speed Rogers, and
Charles F. Walker for not only placing at his disposal the facilities
of the institution, but for suggestions and assistance during the course
of the study. The photographs of preserved specimens were taken
by William L. Brudon, those of habitats in southern Florida by the
senior author and his wife. All of the other illustrations are the work
of the senior author.
We are indebted to Daniel B. Beard, Superintendent, Everglades
National Park, for permission to collect in the area under his jurisdic-
tion. Financial support for field work in southern Florida was fur-
nished by the Museum of Zoology University of Michigan. The
study was completed by the senior author under a grant from the
Bache Fund of the National Academy of Sciences.

AREA OF STUDY
In this study we have concerned ourselves only with the southern
tip of the Florida peninsula and the adjacent Florida Keys. Of course,
reference is made to the area to the north, but the northern boundary
of the present survey consists of an imaginary line from Fort Lauder-
dale to Naples. We have endeavored to examine all specimens from
this area and to visit every accessible part of the region (fig. 1).

PHYSIOGRAPHY AND GEOLOGY
The following brief account of the physiographic and geological
features of southern Florida is based on our personal field experience
and on the works of Bryan and Stoutamire (1930), Cooke (1945),
Henderson (1939), Parker and Cooke (1944), and Schuchert (1929,
1935).


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BULLETIN FLORIDA STATE MUSEUM


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problems. Irving J. Cantrall, Fred G. Thompson, and Harold J.
Walters of the University of Michigan Museum of Zoology, and
H. F. Strohecker of the University of Miami, identified invertebrate
remains from stomach contents. Pierre Dansereau, formerly of the
University of Michigan and now of the Universite de Montreal, greatly
assisted the senior author in formulating the ecological section of this
report; a preliminary paper on this aspect of the study was presented
at the meetings of the American Institute of Biological Sciences and
the American Society of Ichthyologists and Herpetologists in Gaines-
ville, Florida, in 1954.
All of the senior author's work on this project was done at the
University of Michigan Museum of Zoology. Sincere thanks are due
to Norman Hartweg, T. H. Hubbell, the late J. Speed Rogers, and
Charles F. Walker for not only placing at his disposal the facilities
of the institution, but for suggestions and assistance during the course
of the study. The photographs of preserved specimens were taken
by William L. Brudon, those of habitats in southern Florida by the
senior author and his wife. All of the other illustrations are the work
of the senior author.
We are indebted to Daniel B. Beard, Superintendent, Everglades
National Park, for permission to collect in the area under his jurisdic-
tion. Financial support for field work in southern Florida was fur-
nished by the Museum of Zoology University of Michigan. The
study was completed by the senior author under a grant from the
Bache Fund of the National Academy of Sciences.

AREA OF STUDY
In this study we have concerned ourselves only with the southern
tip of the Florida peninsula and the adjacent Florida Keys. Of course,
reference is made to the area to the north, but the northern boundary
of the present survey consists of an imaginary line from Fort Lauder-
dale to Naples. We have endeavored to examine all specimens from
this area and to visit every accessible part of the region (fig. 1).

PHYSIOGRAPHY AND GEOLOGY
The following brief account of the physiographic and geological
features of southern Florida is based on our personal field experience
and on the works of Bryan and Stoutamire (1930), Cooke (1945),
Henderson (1939), Parker and Cooke (1944), and Schuchert (1929,
1935).


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 185


0. ". 6 '. o MARATHON,".3
E G. ( ". RASSY KEY
CUDJOE-'KE'. E T, v "V ,LITTLE PINE C "
CUD" E KE .TORCHE KEY 0 1kKEY VACA
O : -0' "o-- CBAHIA HONDA KEY
STOCK' *** BIG PINE KEY
S ISLAND, RAMROD KEY
SST S S -,y F KEYSUMMERLAND KEY
KE WeST% r SUOGARLOAF KEY
S80BCA CHICA KEY


- 10 MILES


so
I


Figure 1.-Map of southern Florida showing county boundaries, principal
roads, and place names.


The whole of southern Florida, according to most geologists, has
but recently emerged from the sea. The Pamlico beachline of the
Wisconsin now stands at 25 feet above present sea level, and since
no place on the southern Florida mainland has an elevation that
high above sea level, it has been assumed that all of southern Florida
must have been beneath the sea until late Wisconsin time. The re-
lief in southern Florida is slight. Aside from sand dunes on Marco
Island that reach a height of 55 feet above the sea, the highest land
is the east coast ridge or rock rim which averages about 12 feet
above sea level. To the west this ridge slopes fairly abruptly to the


ml


-25


.1


2 2 COLLIER i BROWARD so

NAPLES MILES CITY
S FT LAUDERDALE
W DAVIE*
,o 1 DANIA
DEEP LAKE HOL
j HOLLYWOOD
ROYAL PALM HAM. - ALLAND 26A
C OPE D.-.-.- -- HALLANDALE
MARCO ISLAND ECA E STOWPAND A DADE TA
MONROE STATION PENSUC-O RSIDE
ob St GLADES URFSIDE
4 _SLKEE _. PAOLITA STATION HIALEAH L ON CITY
LOOP RD TAMIAMI TRAIL SWEETWATER MIAMI MIAMI BEACH
S TRAIL CI CREST CORAL GABLES a'-VIRGINIA -KEY
T. IAMIA COCO UT GROVE
PAVILION KEY KENDALL *. -BISCAYNE KEY
CHATHAM R'W 0 MATHESON HAMMOCK
IVf K ; PA ROCKDALEN
SPERRINE-1 UTLER
SGOULOS PETERS
PRINCETON '77
LOSTMANS R.. NARANJA
HOMESTE D SANDS KEY
LON'" ELLIOTT KEY
J;PINE KEY2 X PARA DISE\ ?^if
SHARK R PARADISE
SOUTHERN PENINSULAR FLORIDA V_ KEY

AND THE FLORIDA KEYS CACOpE OT Y R. MADEIRA ABA
COOT BAY RS t
SHOWING LOCALITIES MENTIONED IN THE TEXT S L, /
FLAMING. D o a
D "B ROCK HARBOR
,a o oo TAVERNIER 25

,LO @.-ISLAMORADA
LIGNUMVITAE KYO UPPER MATECUMBE KEY
-INDIAN KEY
LOWER MATECUMBE KEY
LONG KEY
afB' MARATHON









186


BULLETIN FLORIDA STATE MUSEUM


Vol. 3


interior lowlands, which become gradually higher at their western
edge before sloping downward into the Gulf of Mexico. In this low
interior there is also a gradual slope from Lake Okeechobee south-
ward to Florida Bay, so that actually this low region is a broad,
shallow river draining the Okeechobee Basin. The Florida Keys
are also low in elevation, and it is doubtful if any are over 20 feet
above the sea.
Most of the Pleistocene deposits in southern Florida are under-
lain by the Tamiami formation of middle Pliocene age; this formation
is on the surface in the southwestern part of the peninsula and in
small areas between Miami and Fort Lauderdale; in other parts of
southern Florida it underlies the Miami oolite, the Fort Thompson
formation, and the Key Largo limestone. The Tamiami formation is
a calcareous sandstone apparently deposited in the open sea in rather
shallow water far away from land (fig. 2).
The Fort Thompson formation is an alternating freshwater marl
and limestone, and marine shell marl. It contains three separate
marine shell beds possibly corresponding to the Aftonian, Yarmouth,
and Sangamon interglacial periods of the Pleistocene. The interven-
ing freshwater beds were deposited during the Kansan and Illinoian
glacial stages. This formation extends from the south shore of Lake
Okeechobee to central Broward County.
The Key Largo formation, a coral limestone, was formed from a
coral reef developed during the Sangamon interglacial period and
may be slightly older than or contemporaneous with the Miami oolite.
This formation extends from Sands Key on the north to Bahia Honda
Key on the south.
The Miami oolite, an oolitic limestone, also is supposedly of Sanga-
mon age. The formation was deposited in shallow water at a time
when the peninsula north of Lake Okeechobee had emerged, but
all of southern Florida was still under water. The ocean currents
shaped the oolitic ooze into a broad, slightly curving bar that now
forms the eastern rim of the southern part of the peninsula, and that
at one time formed a continuous land mass from Volusia County to
Key West. The Miami o6lite is extensive throughout the southeastern
part of the peninsula and forms the lower keys from, but not includ-
ing, Bahia Honda Key westward. In many places the soft oolitic
limestone is at the surface where it weathers to form pot holes and pits.
The Pamlico sand was deposited during a submergence in Wis-
consin time. A more or less thin sheet of this sand was deposited
over all of southern Florida. In some places it has been eroded away,


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 187

but for the most part it is overlain by recent beach and estuarine de-
posits. The Lake Flirt marl is widely distributed in the Everglades
where it underlies the peaty accumulations and overlies the Pamlico
sand. This marl has accumulated in freshwater lakes probably since
late Wisconsin (post-Pamlico) time and may have continued to do so
locally up to recent time.
Sandy soils derived from the Pamlico sand are prevalent along
the west coast as far south as Marco Island and on the east coast to
Miami. The sandy soils are present on the eastern rim extending
southwestward from Miami. These sands are porous, hold little


-q


Figure 2.-Map of southern Florida showing geological formations and
soil types.


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BULLETIN FLORIDA STATE MUSEUM


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moisture, and are slightly acidic. There are fine-textured sand beaches
along both coasts, but on the keys such beaches are small and local-
ized. The region of the Everglades and associated wet prairies has
a rich peat and muck soil accumulated from the vegetation. This
slightly alkaline soil lies on the Lake Flirt marl and is under water for
considerable periods of time throughout the year. Thin muck de-
posits on the Lake Flirt marl cover most of the western part of the
peninsula. It is on these alkaline soils that the interior pine forests
and cypress forests are developed. The coastal regions not covered
by sand beaches are composed of calcareous marls that are alkaline
in nature. Highly organic soils have accumulated as small islands
in the Everglades and in depressions in the sandy soils. On these
rich soils dense growths of hardwoods (hammocks) are found.

CLIMATE
The climate of southern Florida is an equable one with hot, wet
summers, and mild winters with reduced precipitation. Frost has
been reported occasionally on the mainland, but never on the keys.
The warm winters can be attributed to the Gulf Stream, which
closely follows the east coast, and to the large inland bodies of water,
the Everglades. The average January temperature in Miami is 680
F., and in July it is 81.70 F. Comparative temperatures for Key
West are 69 F. in January and 83.20 F. in July (Mitchell and Ensign,
1928). Rainfall is higher on the mainland than on the keys; for ex-
ample, the average annual rainfall in Miami is 60 inches, on Long
Key 42 inches, and on Key West 38 inches. The greatest amount
of rainfall is during the summer months, culminating with the heaviest
rains in September and October-the hurricane season. Hurricanes
are almost of annual occurrence in some parts of the state. These
tropical storms with winds of high velocity and heavy rains cause
extensive wind damage and flooding. The flooding of the Everglades
makes it necessary for all but the most aquatic animals to seek higher
ground, either the hammocks or the elevated roadbeds.

VEGETATION AND ANIMAL HABITATS
In contrast with the low relief and the equable climatic condi-
tions, southern Florida presents a diversity of environmental condi-
tions. The arrangement and location of vegetation types present in
southern Florida are due primarily to edaphic conditions. The notes
on the vegetation are based on personal experience, and data pre-
sented by Braun (1950), Davis (1943), Egler (1952), Penfound (1952),


_j


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188


BULLETIN FLORIDA STATE MUSEUM


Vol. 3


moisture, and are slightly acidic. There are fine-textured sand beaches
along both coasts, but on the keys such beaches are small and local-
ized. The region of the Everglades and associated wet prairies has
a rich peat and muck soil accumulated from the vegetation. This
slightly alkaline soil lies on the Lake Flirt marl and is under water for
considerable periods of time throughout the year. Thin muck de-
posits on the Lake Flirt marl cover most of the western part of the
peninsula. It is on these alkaline soils that the interior pine forests
and cypress forests are developed. The coastal regions not covered
by sand beaches are composed of calcareous marls that are alkaline
in nature. Highly organic soils have accumulated as small islands
in the Everglades and in depressions in the sandy soils. On these
rich soils dense growths of hardwoods (hammocks) are found.

CLIMATE
The climate of southern Florida is an equable one with hot, wet
summers, and mild winters with reduced precipitation. Frost has
been reported occasionally on the mainland, but never on the keys.
The warm winters can be attributed to the Gulf Stream, which
closely follows the east coast, and to the large inland bodies of water,
the Everglades. The average January temperature in Miami is 680
F., and in July it is 81.70 F. Comparative temperatures for Key
West are 69 F. in January and 83.20 F. in July (Mitchell and Ensign,
1928). Rainfall is higher on the mainland than on the keys; for ex-
ample, the average annual rainfall in Miami is 60 inches, on Long
Key 42 inches, and on Key West 38 inches. The greatest amount
of rainfall is during the summer months, culminating with the heaviest
rains in September and October-the hurricane season. Hurricanes
are almost of annual occurrence in some parts of the state. These
tropical storms with winds of high velocity and heavy rains cause
extensive wind damage and flooding. The flooding of the Everglades
makes it necessary for all but the most aquatic animals to seek higher
ground, either the hammocks or the elevated roadbeds.

VEGETATION AND ANIMAL HABITATS
In contrast with the low relief and the equable climatic condi-
tions, southern Florida presents a diversity of environmental condi-
tions. The arrangement and location of vegetation types present in
southern Florida are due primarily to edaphic conditions. The notes
on the vegetation are based on personal experience, and data pre-
sented by Braun (1950), Davis (1943), Egler (1952), Penfound (1952),


_j


II









1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 189

Safford (1919), and Small (1931a,b,c, 1933). The nature of the vegeta-
tion and composition of the flora are discussed under the descriptions
of the various ecological situations. The classification of habitats
used here is a somewhat modified and simplified version of that pro-
posed by Davis (1943). The following classification of habitats (with
the exception of VI) consists of three levels: those indicated by Roman
numerals differ in the amount of moisture present; those indicated
by capital letters differ in the structure of the vegetation; those in-
dicated by Arabic numbers differ floristically. The classification of
animal habitats in southern Florida is outlined as follows: 2


I. XERIC
A. Sandy scrub
B. Pine forest
1. Sandy pineland
2. Rocky pineland
C. Beaches
II. MESIC
A. Hammocks
1. Oak
2. Cabbage palm
3. Tropical
III. ALTERNOHYGRIC
A. Cypress flats
B. Prairies
1. Saw grass
2. Poverty grass
3. Needle grass


IV. HYGRIC
A. Cypress heads
B. Sink ponds
C. Canals and rivers


V. HALOHYGRIC
A. Salt marshes
B. Mangrove swamps


VI. EDIFICARIAN AND
RUDERAL


In the following pages the various habitats as proposed above
are discussed, giving their distribution in southern Florida, soil con-
ditions when these are known, structure of the vegetation, and a
list of the dominant or more common plant species. Figure 3 shows
the rough distribution of the generalized habitats.
I. XERIC
Dry conditions are quite common in southern Florida. The east-
ern rock rim made up of an oolitic limestone and overlain by a thin

The terms used here to denote the major realms of ecological division do
not correspond with similar terms derived by del Villar (1929).
The term "alternohygric" is proposed for areas in which there is alternate
flooding and drying out, such as is typical of the Everglades.


Ii











BULLETIN FLORIDA STATE MUSEUM


Vol. 3


cover of sand is typical of the xeric conditions. This situation also
is present on the lower keys where the soil conditions are much the
same. The sandy flatlands north of Miami and north of Naples, plus
the greater part of Marco Island are remnants of Pleistocene beaches.
Porous sandy soils that hold little moisture apparently are the con-
trolling factors in determining the seemingly arid conditions. Pines
of one species or another and the palmetto, Serenoa repens, are com-
mon to all xeric habitats in Florida.
A. SANDY SCRUB.-This is the Pinus clausa-Ceratiola ericoides as-
sociation commonly referred to as the rosemary scrub. This associa-


.': : -" : + :" :7" .7 -,-_-:-__-_ --------=-- 3:-- --- '


IV
1/ -. - ..

.s
SpL




-~-v-
-- z

] SCei L
: __=.. .-- -- _----__.--- : .
', : .-t--*- -- - -- "


] FlUEI

]: -% --.-
R'i


--'YPREF4


]


PMANGRilE
MANGROvE


-5 --- SALINITY BORDER


~ hn'. -'-'
3 d '**
/ :"'


Figure 3.-Map of southern Florida showing general distribution of major
types of vegetation.


_I


K
U


I'-- I-----------~


I
e 2




-26


25'


liP' ~
i -
r
-a" '




El


1958 DUELLMAN AND SCIIWARTZ: AMPHIBIANS AND REPTILES 191

tion is found on sandy flatlands. In southern Florida it is restricted
to a few small patches between Miami and Ft. Lauderdale on the
east coast and to scattered patches in the vicinity of Naples and Marco
Island on the Gulf coast. In this habitat the pines are widely spaced,
often 100 yards apart. Ceratiola ericoides, Quercus myrtifolia, and
Serenoa repens comprise the only real cover. Sometimes these plants
form dense thickets covering many square yards to a height of four
or five feet. Opuntia is also found in this association; graminoids
are local (fig. 4).
B. PINE FoREsT.-Pine forests are found on the sandy soils along
the east coast and Gulf coast, on the eastern rim extending southwest-
ward from Miami, on the lower keys, and on the sandy and marly
soils of the interior in the western part of the peninsula.
1. Sandy pineland.-The pine flatwoods are rather extensive in
the sandy regions north of Miami and in the western part of the
peninsula north and east of Naples. In these areas the pine forests
have developed on ground-water podzols, such as the Pamlico sand.
The pine flatwoods in the interior have developed on soils with a
calcareous substratum. In the former, Pinus caribaea is the dom-
inant tree (fig. 5), whereas in the latter, Sabal palmetto often com-
bines with the pines to form the uppermost story, and in some places
the cypress, Taxodium ascendens, intermingles with the pines (fig. 10).
Often the forests are rather open, especially those in the interior
where, as often as not, the structure of the vegetation more closely
approaches that of a savanna than a forest. The understory consists
of small shrubs, such as Vaccinium myrsinites, Ilex glabra, Quercus
pumila, and Pycnothymus, aside from the omnipresent Serenoa repens.
Typical graminoids in this association are Andropogon and Aristida.
2. Rocky pineland.-The pineland on the eastern rock rim
formed of Miami oolite may be considered as being distinct from the
sandy pineland, not in regard to the pine, but to the associated plants.
The rocky pineland is nearly continuous from Ft. Lauderdale to
Miami and southwest to Long Pine Key in the midst of the Everglades.
The pine forests are well developed on the lower keys, especially Big
Pine Key. These are open pine forests growing on an oolitic lime-
stone covered with a thin layer of sandy soil. In many places there
is no soil, so that the eroded limestone stands at the surface. The
dominant plant is Pinus caribaea (fig. 6). Other trees, such as Coc-
cothrinax argentea, Metopium toxiferum, and Guettarda scabra, are
found scattered among the pines. The common grasses are Andropo-
gon and Aristida stricta. The pine forests on the lower keys are much


h




I


'-7-~--~~e~a~lsaa lr


Rguge 4-Rmm-y-crb habitat -n M-ro bl,,d, CUi,, C-tyh. I.
'h f..g...ud C-H.I Hericid- ..nd Q.-csmrtfln Th ft. Ui Pi..s





clnL am










1958 DUELLMAN AND SCHWART-L AMPHIBIANS AND REPTILES 193


Figr 7.-B-1, -o th, --wad sid, f Mu- sladCob-e C ...ty.
C-h I, i th dai ... t pl,.t I, th f-g ... d; _h th b.,kg ... d i U.0N.o









BULLETIN FLORIDA STATE MUSEUM


Vol. 3


the same as those on the mainland, but usually less dense. Also,
Cocos nucifera has been introduced into the pinelands, where it forms
a dominant part of the vegetation in some areas. Mesophytic ham-
mock invades the pineland in many areas and soon becomes dominant.
In southern Florida the Pinus caribaea-Serenoa repens association
apparently is a fire disclimax, for in areas where the pine forests have
been protected from fire, hammock vegetation is increasing to the
detriment of the pine.
C. BEACHEs.-Extensive fine-textured sand beaches are found
from Ft. Lauderdale to Miami Beach on the east coast and in the
vicinity of Naples and on Marco Island on the Gulf coast. Small
sandy beaches are found on some of the Florida Keys. The principal
plants include Coccoloba uvifera, Uniola, and Cenchrus (fig. 7). Cocos
nucifera has been planted on the inland side of many beaches.

II. MESIC
The existence of mesic habitats in southern Florida is usually due
to pockets in the marl or rock substratum in which organic soils have
accumulated. Consequently, the mesic habitats are in the form of
"islands" scattered throughout the Everglades and pinelands.
A. HAMMOCKS.-The term hammock in southern Florida is ap-
plied to patches of woodlands, other than pine. Hammocks may be
high and relatively dry, such as oak hammocks; fairly well drained
but subject to wet conditions at least during parts of the year, such
as cabbage palm hammocks; or they may be in the form of a dense
growth of subtropical and tropical plants that form an actual jungle,
such as the tropical hammocks.
1. Oak hammock.-This type of mesic habitat is restricted to a
few places on the eastern rim south of Miami, in Coral Gables, and
in South Miami. Quercus virginiana is the dominant tree; Ficus
aurea and F. brevifolia are commonly found. Other plants associated
with the oak hammocks include Coccoloba laurifolia, Metopium toxi-
ferum, and Psychotria; Tillandsia is everywhere present in the trees.
2. Cabbage palm hammock.-This type of hammock is most com-
monly found along the western margin of the prairies and scattered
through the cypress flats and pine woods in the western part of the
peninsula. In these areas the hammocks are formed on marl and muck
soils. Sabal palmetto is the dominant plant, and in hammocks where
the trees are not too dense there is an understory that may include
Serenoa repens, Yucca aloifolia, and many species of small herbs
(fig. 8).


El_


II





































of Serene rpen near Cretown, Coler County


088ty' AZ-8g 818 pRk-,8 88 he, pict-88 888 N888,88109 mltt, E88888
Th88sE, Km8d,,d8, PsR8888s, P-W80, -d B.rss8-.


1958 DUELLMAN AND SCHWAR17: AMPHIBIANS AND REPTILES 195









BULLETIN FLORIDA STATE MUSEUM


3. Tropical hammock.-These hammocks may be rather exten-
sive, such as Matheson Hammock south of Coral Gables, Royal Palm
Hammock, Paradise Key, and those along the southern coast and on
Key Largo, or they may be small clumps of trees, such as most of
the hammocks in the Everglades. These "tree islands" in the wet
prairies are most common along the eastern margin of the Everglades,
especially south of the Tamiami Trail. In the Everglades they have
formed on ground slightly higher than the surrounding prairies; on
the rock rim they are more often found in depressions in the rock
where organic material has accumulated and provided a place for
the luxuriant growth of trees. The floor of the hammocks is usually
nearly devoid of herbaceous plants, with the exception of ferns. In-
stead the ground is covered with leaves and leaf mold. It is in these
hammocks that the West Indian elements in the flora are most ap-
parent, there being scores of plant species with Bahaman and Cuban
affinities. A complete list of the more common and dominant plants
is not feasible here, but some of the most frequently encountered and
most noticeable species in the hammocks on the mainland are listed
below.
Bursera simaruba Metopium toxiferum
Coccoloba laurifolia Psychotria
Dipholis Roystonea regia
Eugenia Sideroxylon foctidissimum
Ficus area Swietenia mahogani
Lysiloma bahamensis Tillandsia
Royal Palm Hammock in Collier County is a fine example of a
Roystonea hammock; the large palms tower above the other vegeta-
tion. Roystonea and Ficus are dominant trees on Paradise Key (fig.
9). Southwest of Paradise Key is a hammock composed almost en-
tirely of Swietenia mahogani. The plants comprising the hammocks
on the mainland are also found in hammocks on the keys; however,
more West Indian species are encountered on the keys. Swietenia
mahogani is more common on the keys, while Thrinax, Pseudothrinax
sargentii, Guaiacum sanctum, Hippomane mancinella, and Piscidia
communis also are commonly found. The hammocks on the keys do
not attain the luxuriant splendor of those on the mainland; the trees
are more crowded and stunted, a condition that may be the direct
result of the tropical storms that batter the islands rather frequently.

III. ALTERNOHYGRIC
The term alternohygric is applied to those environments in south-
ern Florida that are subject to periodic flooding and drying. More


____ i~___~


__ ____


Vol. 3











1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 197


Fig- 1 0--Cyp- f fit 47 ike -, f hfim, M-no C.-nty. H-r
Tordmiu m .. dmn mi.. withU Pim sconb-o









BULLETIN FLORIDA STATE MUSEUM


specifically, it is applied to the low flat interior region of the southern
part of the peninsula, that region in which the wet prairies and cy-
press flats are found. The soils are either a combination of peat and
muck or marl and muck.
A. CYPRESS FLATs.-The cypress savannas are extensive in the
western part of the peninsula where they are interrupted by expanses
of pineland and wet prairies. Also, scattered through the cypress
savannas are tropical and cabbage palm hammocks. Taxodium ascend-
ens is the only tree distributed throughout the habitat (fig. 10). Serenoa
repens is plentiful and in certain areas forms a dense understory. The
ground cover consists primarily of Utricularia juncea, Cyperus Haspan,
and Rynchospora.
B. PRAIRIES.-The flat grasslands of southern Florida form the
most extensive vegetational formation, and it is this plant formation
that has received the name "Everglades." These grasslands extend
from Lake Okeechobee to Cape Sable. For the most part they are
formed on peat and muck soils, and in spite of the highly organic
composition of the soils the underlying limestone creates a slightly
alkaline condition.
1. Saw grass.-The prairies that are made up almost entirely of
Mariscus jamaicensis are the true Everglades. They lie in the middle
of the prairie region, being bordered on each side by the poverty-
grass prairies. To the south they fuse with the brackish salt marshes
or mangrove swamps. The saw grass prairies are homogeneous. Mar-
iscus is by far the most common plant, while Peltandra virginica, Soli-
dago fistulosa, and Polygonum occur in scattered areas in much fewer
numbers (fig. 11). The continuity of the saw grass marshes is inter-
rupted only by hammocks, ponds, and canals. Open ponds in the
Everglades, locally called sloughs, are usually bordered by dense
growths of willow, Salix amphibia (fig. 12).
2. Poverty grass.-The prairies immediately bordering the higher
ground-the eastern rim and the sandy flatlands to the west-are
rather different floristically from the true Everglades or saw-grass prai-
ries. In the bordering areas Mariscus is scarce, the vegetation being
composed almost entirely of Andropogon and Aristida. A possible
explanation for the floristic differences may lie in the slightly different
soil conditions towards the borders of the interior lowlands; possibly
Mariscus is able to grow only in the richest soils, these being present
only in the middle of the prairie region.
3. Needle grass.-The needle-grass prairies composed almost en-
tirely of Eleocharis cellulosa are found in the southeastern part of


II


Vol. 3










1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 199
























FRgoon 12-Taylor Slogb t P .. dme Koy, Dade County. Tho p,,d is
,nm-dod by Sd on,,phibin nod Typha.
























Fig", 13.-N,,dA 9g- po 999, oomposd of El .. lou celllosa, ut th,
.B9nit b i i
C ...ty.
abow, th, lono-k in th, distto- mr p~lon, P ... ti,









200 BULLETIN FLORIDA STATE MUSEUM Vol. 3

the peninsula, south and east of the rock rim that extends southwest-
ward from Miami. Here the soils are more calcareous than those in
the interior lowlands, and there is a much greater salinity effect, which
is especially prominent near the coast where Distichlis and Spartina
are found with the Eleocharis. The salinity border may be determined
by the presence of the tall slender palms, Paurotis (fig. 13).

IV. HYGcIC
The strictly aquatic freshwater habitats in southern Florida are
found naturally only in cypress ponds, sink ponds, and in the few
real rivers. The most extensive freshwater habitat, however, is in
the form of artificial canals, most of which parallel the roads in the
area.
A. CYPRESS HEADS.-The so-called cypress heads or ponds are de-
pressions that contain water throughout the year and support a more
or less dense growth of cypress. These situations are most frequently
met with west of the Everglades, but some cypress heads occur in the
Everglades proper. Both Taxodium distichum and Taxodium ascend-
ens make up the dominant vegetation in these areas (fig. 14). Associ-
ated plants are Magnolia virginiana, Salix amphibia, Isnardia repens,
Sagittaria, and Thalia geniculata.
B. SINK PONDS.-Ponds that are the result of solution of the lime-
stone substratum are common in central Florida, a region of typical
karst topography. There are few such ponds in southern Florida;
one of the largest is Deep Lake in Collier County. There are several
small sinks and solution pits on the eastern rim and one large pit on
Big Pine Key (fig. 15). Nymphaea and Typha are typical plants of the
shores of these ponds.
C. CANALS AND RIVERS.-The network of canals in southern Florida
that were built for drainage purposes form an important and exten-
sive habitat that contains water throughout the year. The introduced
water hyacinth, Eichhornia (Piaropus) crassipes is the dominant plant
in the canals. In many places these hyacinths have become so thick
that they form a solid floating mass on the surface of the water. Typha,
Pontederia, Sagittaria, and Nymphaea also contribute to the canal veg-
etation. Salix amphibia and Taxodium are common canal-edge plants.
Along the Tamiami Trail and some other roadways in southern Florida,
Casuarina equisetifolia has been planted between the roads and the
canals (fig. 16). The elevated roadbeds paralleling the canals offer
a site of refuge for many animals when the prairies are flooded.


---------I --~Ii




































Figure 14.--Cyp-ss p,,d 5.5 mil,,*-th d M-- Stb,, Coli,, C ..


Figr 15.-oltU,, t mn Big n.e K- M.na C..ty/. Th, wga-ho
_...uding th sink i -p-ord of Pi.,. foib- ..ad S-.- n rep-s


1958 DUELLMAN AND SCHWARTZ AMPHIBIANS AND REPTILES 201










202 BULLETIN FLORIDA STATE MUSEUM V.I. S


2,-


Fig- 16.-Th. Taooi C .. 20 milm -t of Mind, Skd, Cooy,.
O. th, Aght book ... Typha ood SohM,.mphibi,. O, theUIA m pl,,ted C-
42,2 oq,22SifoliS.


'---








1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 203

There are few real rivers in southern Florida. The water hyacinth
usually is absent from the rivers; Taxodium and Salix are usually pres-
ent along the banks; also, the banks may be lined in certain areas
with tropical hammock (fig. 17).
V. HALOHYCGIC
Because there is so little relief in southern Florida, tides, storms,
and rainfall cause a fluctuation of the salinity border. In the southern
part of the Everglades there is a gradual transition between fresh-
water plant species and those commonly associated with brackish
conditions.
A. SALT MARSHES.-In the southwestern part of the state brackish
marshes or prairies are an intermediate stage between the freshwater
prairies and the coastal mangrove swamps. These are formed on cal-
careous soils. The most extensive salt marsh is on Cape Sable (fig. 18).
The following herbaceous plants are common in this habitat: Distichlis
spicata, Spartina, Juncus Roemerianus, Batis, and Salicornia.
B. MANGROVE SWAMPS.-The dense tangle of mangrove swamp
fringing the southern edge of the Florida peninsula and the Florida
Keys is one of the most extensive in the world. The low southwestern
coastal region, the Ten Thousand Islands, and the smaller keys have
practically no vegetation other than mangroves, which flourish on
the coastal marl. The mangroves show a definite order of succession.
Rhizophora mangle is farthest from shore; then follows Avicenna
nitida, Laguncularia racemosa, and finally Conocarpus erecta (fig. 19).
Batis, Salicornia, and Spartina are found among the mangroves.
VI. EDIFICARIAN AND RUDERAL
The man-made habitats form an important factor in the distribu-
tion of many species of reptiles and amphibians in southern Florida,
especially those introduced into port cities from the West Indies.
Edificarian and ruderal situations are frequently inhabited by several
native species as well. In many places shaded gardens and yards
provide the only moist habitats available to some populations of am-
phibians; citrus groves and cultivated fields likewise form habitats
for many species.

COMPOSITION OF THE HERPETOFAUNA
In the southern part of the Florida peninsula and on the Keys
are found 80 species of amphibians and reptiles, not including five
species of sea turtles. Of the 80 species, eight were undoubtedly in-
troduced into the area, and four others may have been introduced.4


I








Vol. 3


kiggr 0B.Sat mmrh mn Cap, Sabie, M---o C-uty. Dis~ahlk, Spanim,
S'If ... l, aud Bati, u, pmelmt , the .,,h. 1, th, b,,kgraud i~ataagla f
...g,.-es


rr~ka~[r~i~L~PB~"~;~BB


204 BULLETIN FLORIDA STATE MUSEUM


.,
:o y
1~4 -~I .I
.C


crwrr~, v rw.~k.









1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 205

Thus, only 68 native species may exist. These comprise 51 genera
(amphibians 12, reptiles 39), and 21 families. For a warm lowland
area this is a relatively small number of species. However, an im-
poverished herpetofauna is what might be expected at the end of
a long peninsula, through the length of which certain habitats and
their inhabitants disappear. The depauperate nature of the herpeto-
fauna is well illustrated by comparing the numbers of families, genera,
and species of amphibians and reptiles known from northcentral
Florida with those occurring in southern Florida. In the north are
24 families, 62 genera, and 101 species. The Ambystomidae, Pletho-
dontidae, and Amphisbaenidae do not reach southern Florida. In
the latter area are four genera and species of salamanders, as com-
pared with ten genera and 15 species in northcentral Florida. Croco-
dylus acutus is the only species (except the introduced ones) in
southern Florida that does not extend to the northern part of the pen-
insula. In comparison with the above figures, in all of the eastern
United States there are 26 families, 75 genera, and 186 species. Al-
though the southeastern states generally have a greater number of
species of amphibians and reptiles than do the northern states, as one
moves southward in peninsular Florida the number diminishes.
For obvious reasons the five sea turtles-Chelonia mydas, Eretmo-
chelys imbricata, Lepidochelys olivavcea, Caretta caretta, and Dermo-
chelys coriacea-that frequent the coastal regions of the southeastern
United States are not included in the above analysis.
Nine lizards and three anurans that have affinities with faunas
other than that of North America are found in southern Folrida. Seven
lizards and two frogs have West Indian affinities: Gonatodes fuscus,
Sphaerodactylus argus, S. cinereus, S. notatus, Anolis distichus, A.
sagrei, Leiocephalus carinatus, Eleutherodactylus ricordi, and Hyla
septentrionalis. Of the above species, Leiocephalus carinatus and
Sphaerodactylus argus apparently are no longer extant in southern
Florida. Of the other nonnative forms, Bufo marinus may have been
introduced from almost anywhere around the Caribbean or Gulf of
Mexico; Ameiva ameiva is from South America, and Hemidactylus
turcicus occurs in tropical regions around the world. Four of the
above species are rather widely distributed in southern Florida and
inhabit natural, unmodified areas; these are Eleutherodactylus ricordi,
Hyla septentrionalis, Sphaerodactylus notatus, and Anolis sagrei. Ei-
ther these species have inhabited southern Florida for a much longer

4Neill (1957: 207) mentioned the presence of Anolis equestris in southern
Florida, but did not give a locality for the species. It is not included here.


U.


i*I









BULLETIN FLORIDA STATE MUSEUM


time than the others or they have certain characteristics better adapt-
ing them for life in southern Florida. Anolis sagrei possesses morpho-
logical differences that distinguish it from the West Indian popula-
tions. It therefore seems logical to assume that A. sagrei has been
in Florida for a considerable length of time. The other eight species
for the most part, with the exception of E. ricordi, are found only in
port cities and appear to be recent introductions through the agency
of man. Crocodylus may have reached southern Florida by way of
the West Indies.

ECOLOGY OF THE HERPETOFAUNA
As has been shown in the section on vegetation, the area studied
may be divided into five major natural habitats. Certain factors tend
to impair the accurate specification of the herpetological assemblage
characteristic of each of these habitats, and also the distribution of
these assemblages.
First, there is the element of personal error, that is, the lack of
uniformity in the study of the several habitats. Because of their in-
accessability many large but extensive tropical hammocks were not
studied. The hordes of mosquitoes tested the tolerance of collectors
in mangrove swamps; a minimum of time was spent there. In areas
where collecting was poor, we often gave up in despair and departed
for more productive localities. In some of the areas mentioned we
undoubtedly missed certain species.
Second, the seasonal activity of the fauna, particularly the anurans,
precludes a complete faunal picture. Although all areas in southern
Florida were studied in the wet summers and in the dry winters, it
was not always possible to be at the right place at the right time.
The element of chance figured strongly the night we first heard
Hyla gratiosa in southern Florida.
Third, extensive cutting, burning, draining, and other human
forms of destruction unquestionably have altered patterns of distribu-
tion, possibly restricting the ranges of some species and extending
those of others. This factor raises several questions: What were the
distributions of aquatic and semiaquatic species prior to the building of
canals? What were the distributions of species now found in the Ever-
glades before the roadways were built and large sections of the prai-
ries drained? These questions can never be answered; we can only
hope to ascertain the present distributions on the basis of the habitats
now in existence.
Finally, the principal criteria used by zoogeographers are the pres-
ence and absence of species. However, in some instances the rela-


-- ---


Vol. 3










1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 207

tive abundance of some or all of the species may be as indicative
of faunal similarities and differences as is the presence. We have
attempted, in a rather crude fashion, to determine relative abundance,
that is, noting whether a species was of apparently rare, moderate,
or abundant occurrence in a given habitat.
In presenting the following picture of the local distribution of
the herpetofauna in southern Florida, we have attempted to weigh
our data and observations with full knowledge of the sources of
possible error. In some instances our interpretations have been
based more on our observations of the species outside of the area of

TABLES 1 and 2

DISTRIBUTION OF NONMARINE REPTILES AND AMPHIBIANS OF SOUTHERN FLORIDA

2. Distribution
1. Distribution by by geographic
major habitats regions

Species M Q





Siren lacertina M A X -
Pseudobranchus s. belli M A X -
Notophthalmus v. piaropicola R M A X -
Amphiuma means M A X -

Scaphiopus holbrooki A R R X X
Bufo marinus -R X -
Bufo quercicus M R A M X X
Bufo t. terrestris A M M M X X
Eleutherodactylus r. planirostris M A M X X X
Acris g. dorsalis A X -
Pseudacris n. verrucosa M A X -
Limnaoedus ocularis A X -
Hyla cinerea M M A X X X
Hyla femoralis A R X
Hyla gratiosa A R X -
Hyla septentrionalis A M X X X
Hyla squirella M M A M X X X
Gastrophryne carolinensis M M A R X X X
Rana a. aesopus R -
Rana grylio M A X -
Rana pipiens M A M X X




UT-~ _________


208 BULLETIN FLORIDA STATE MUSEUM Vol. 3

TABLES 1 and 2
DISTRIBUTION OF NONMARINE REPTILES AND AMPHIBIANS OF SOUTHERN FLORIDA

2. Distribution
1. Distribution by by geographic
major habitats regions



o




Chelydra osceola M A X
Sternotherus odoratus M A -
Kinosternon b. bauri A M - X
Kinosternon b. palmarum M A X X -
Kinosternon s. steindachneri M A R X -
Terrapene c. bauri A M M X X X
Malaclemys t. macrospilota A X -
Malaclemys t. rhizophorarum A X X
Malaclemys t. tequesta A X -
Pseudemys f. peninsularis M A X -
Pseudemys nelsoni M A X -
Deirochelys r. chrysea M X
Gopherus polyphemus R X
Trionyx ferox A X -

Crocodylus acutus M X X X
Alligator mississipiensis M A M X X X

Gonatodes fuscus A X
Hemidactylus turcicus A X X
Sphaerodactylus a. argus ? ?
Sphaerodactylus cinereus A X
Sphaerodactylus notatus M A M X X X
Anolis carolinensis M A M M X X X
Anolis distichus A M X -
Anolis s. stejnegeri M A X X
Leiocephalus c. virescens ? ? -
Sceloporus woodi A X -
Ophisaurus a. longicaudus R X -
Ophisaurus compressus A M -X
Ophisaurus ventralis A M X -
Ameiva ameiva M -X -
Cnemidophorus s. sexlineatus A M X X X
Lygosoma laterale M A -M X X X
Eumeces e. egregius A M X X
Eumeces e. onocrepis M - X -










1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 209

TABLES 1 and 2
DISTRIBUTION OF NONMARINE REPTILES AND AMPHIBIANS OF SOUTHERN FLORIDA

2. Distribution
1. Distribution by by geographic
major habitats regions

Species a





Eumeces inexpectatus M A R M X X X
Eumeces laticeps R X -
Neoseps reynoldsi R X -

Diadophis p. punctatus M A R X X X
Heterodon platyrhinos M X X -
Farancia a. abacura A M X -
Coluber c. paludicola M A X X -
Coluber c. priapus A M X X
Masticophis f. flagellum A X -
Opheodrys aestivus M M A X X X
Drymarchon c. couperi M A M X X X
Elaphe g. guttata A M R M X X X
Elaphe o. quadrivittata M A X X ?
Elaphe o. rossalleni M A X -
Pituophis m. mugitus R X -
Lampropeltis d. doliata A M X ?
Lampropeltis g. floridana M M A X -
Cemophora coccinea M M X -
Tantilla c. wagneri A M X X
Natrix c. floridana M A X -
Natrix s. compressicauda A X X X
Natrix s. pictiventris M A X -
Natrix taxispilota R M X -
Seminatrix p. cyclas M X -
Storeria d. victa A M X X
Thamnophis s. sackeni M R A X X
Thamnophis s. sirtalis M R M X -
Liodytes alleni A X -
Micrurus f. fulvius M A X X -
Agkistrodon p. piscivorus M A M X X X
Sistrurus m. barbouri R A X -
Crotalus adamanteus A R X X X


A, abundundt; M, moderately abundant; R, apparently rare; X, present; ?,
presence not established, or if formerly present now apparently extinct.
Thamnophis~ ~ ~ ~ ~ ~ 1.sceiIM









BULLETIN FLORIDA STATE MUSEUM


study and on the work of others than on the data derived from our
studies in southern Florida.
Since in its broader aspects vegetation is a manifestation of the
combined effects of climate, substrate, and topography, all of which
in large measure determine animal distributions, it naturally follows
that a high degree of coincidence between faunal and floral bound-
aries is to be expected. An analysis of the herpetofauna bears out this
supposition. It is true that in some instances, aquatic turtles for ex-
ample, this correlation may break down. In table 1 the distribution
of the amphibians and reptiles of southern Florida is given for the
five major natural habitats plus "edificarian and ruderal." Further-
more, notation is made as to the apparent abundance of the individual
species in each habitat.
The principal division of habitats is between the xeric (pine forests
and scrub) and the alternohygric (prairies and cypress flats). Twenty-
two species occur in the xeric habitats but not in the alternohygric
ones; these include:
Scaphiopus holbrooki Lygosoma laterale
Eleutherodactylus ricordi Eumeces egregius
Hyla femoralis Neoseps reynoldsi
Hyla gratiosa Heterodon platyrhinos
Rana areolata Masticophis flagellum
Gopherus polyphemus Pituophis melanoleucus
Sphaerodactylus notatus Lampropeltis doliata
Anolis sagrei Cemophora coccinea
Sceloporus woodi Tantilla coronata
Ophisaurus attenuatus Micrurus fulvius
Cnemidophorus sexlineatus Crotalus adamanteus
Although Hyla femoralis and Hyla gratiosa have been found in
cypress flats, their principal distribution appears to be in the pine
forests and scrub habitats; thus, they are included in the list. Of
the above species, Gopherus polyphemus, Sceloporus woodi, and
Neoseps reynoldsi appear to be restricted, at least in southern Florida,
to the rosemary-scrub habitat. On the other hand, Masticophis flagel-
lum and Pituophis melanoleucus have been found only in pine forests
in southern Florida. Heterodon platyrhinos has been encountered
in pine forest and in scrub. Anolis sagrei most commonly inhabits
edificarian situations, but where it occurs in natural situations, it is
found in xeric pine forest. The other 11 species are found in xeric
habitats and mesic hammocks.
Twenty-one species inhabit the wet pariries, cypress flats, and
aquatic situations in these environments, but do not range into the
pine forests, scrub, or hammocks. This group includes:


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 211


Siren lacertina
Pseudobranchus striatus
Notophthalmus viridescens
Amphiuma means
Acris gryllus
Limnaoedus ocularis
Rana grylio
Chelydra osceola
Sternotherus odoratus
Kinosternon bauri
Kinosternon subrubrum


Pseudemys floridana
Pseudemys nelsoni
Alligator mississipiensis
Farancia abacura
Natrix cyclopion
Natrix sipedon
Natrix taxispilota
Seminatrix pygaea
Liodytes alleni
Agkistrodon piscivorus


Of these, Acris gryllus, Limnoaedus ocularis, Farancia abacura,
Liodytes alleni, and Agkistrodon piscivorus are inhabitants of the
prairies and flats, but do not display essentially aquatic habits. Pseud-
obranchus striatus, Notophthalmus viridescens, small Siren and Amph-
iuma, and Seminatrix pygaea live in the masses of aquatic vegetation,
but not in open water. The other species are more or less aquatic.
Efts of Notophthalmus viridescens have been found in hammocks,
but so far as is known the adults inhabit only the aquatic situations
in the prairies and cypress ponds. Other common inhabitants of
the prairies, but not restricted to this habitat include:


Bufo quercicus
Hyla cinerea
Hyla squirella
Rana pipiens
Coluber constrictor


Elaphe obsoleta
Lampropeltis getulus
Thamnophis sauritus
Sistrurus miliarius


Although 34 species have been encountered in one or more kinds
of hammocks in southern Florida, and 11 species appear to reach
their greatest abundance in this mesic habitat, it is doubtful if any
are restricted to this environment. Eumeces laticeps, a little-known
species in southern Florida, has been collected only in oak hammock.
Hyla septentrionalis and Anolis distichus, both introduced species that
occur in edificarian situations, are most abundant in mesic habitats.
Eleven species occur in pine forests and in hammocks, but not in the
prairies. Eighteen species are common to all three habitats, namely:


Bufo quercicus
Bufo terrestris
Hyla cinerea
Hyla squirella
Gastrophryne carolinensis
Terrapene carolina
Anolis carolinensis
Eumeces inexpectatus
Diadophis punctatus


Coluber constrictor
Opheodrys aestivus
Drymarchon corais
Elaphe guttata
Elaphe obsoleta
Lampropeltis getulus
Storeria dekayi
Thamnophis sauritus
Thamnophis sirtalis










BULLETIN FLORIDA STATE MUSEUM


The above species are among the most abundant and widespread
amphibians and reptiles in southern Florida; of these, only Anolis
carolinensis, Eumeces inexpectatus, Diadophis punctatus, and Dry-
marchon corais appear to reach their greatest abundance in the mesic
habitats.
Seven species are known to occur in the pine forests or rosemary
scrub and in the wet prairies or cypress flats, but not in the mesic
hammocks; these are:
Pseudacris nigrita Ophisaurus compressus
Hyla femoralis Ophisaurus ventralis
Hyla gratiosa Sistrurus miliarius
Rana pipiens
As stated previously, Hyla femoralis and Hyla gratiosa are essen-
tially restricted to the pine and scrub habitats; they enter the cypress
area where pine and cypress mix and interdigitate in central Collier
County. Sistrurus and Rana pipiens, for the most part, are inhabi-
tants of the prairies, but where prairie and pine habitats meet they
are often encountered in the pine woods. Both species of Ophisaurus
are most frequently encountered in grassy areas, whether these be
in the pine savannas or in the prairies. Pseudacris nigrita appears to
be associated with the surface-pitted limestone, a condition that ap-
pears frequently in the prairies and in the pine woods. This species
is particularly abundant in the pine-prairie ecotone in Dade County.
The aquatic turtles Deirochelys and Trionyx, although inhabiting
permanent water in the Everglades and associated habitats, appar-
ently do not range into the real prairies. Crocodylus, the three forms
of Malaclemys terrapin, and Natrix sipedon compressicauda, together
with the 5 species of sea turtles, are confined to salt- or brackish-
water habitats. Kinosternon bauri, Alligator mississipiensis, and Ag-
kistrodon piscivorus are frequently found in salt or brackish water.
Restricted to edificarian situations are 3 species of lizards, all
of which have the earmarks of rather recent introductions-Gona-
todes fuscus, Hemidactylus turcicus, and Sphaerodactylus cinereus.
Other introduced forms along with certain native species are found
in greater or lesser abundance in the edificarian-ruderal habitat.
From the above analysis it is clear that although none of the
major habitats has a highly restricted herpetofauna, each has charac-
teristic species. Forty-five species inhabit the xeric habitats, and 7
of these are confined to that habitat; 34 inhabit the mesic, and 1 is
confined; 46 inhabit the alternohygric, and 4 are confined; 21 inhabit
fresh water, and 2 are confined; 7 inhabit brackish water, and 3 are


__


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 213

confined (sea turtles not included); 18 inhabit edificarian-ruderal sit-
uations, and 3 are confined. Not included are Ameiva ameiva and
Bufo marinus, both recently introduced species, and Leiocephalus
carinatus and Sphaerodactylus argus, both of which were introduced
into edificarian habitats but apparently did not survive.
In three different instances two races of a species inhabit different
environments in southern Florida. Coluber constrictor paludicola
ranges throughout the prairie habitats and sometimes enters the mesic
hammocks; C. c. priapus inhabits the pine forests, scrub, and ham-
mocks. Elaphe obsoleta has a similar distribution, with the race
quadrivittata being found in the pine forests and hammocks, whereas
the race rossalleni is essentially confined to the prairies. In Natrix
sipedon, the subspecies pictiventris inhabits freshwater ponds and
streams, the prairie, and cypress flats; the subspecies compressicauda
is restricted to the brackish mangrove swamps and salt marshes.

GEOGRAPHY OF THE HERPETOFAUNA
In order to gain an adequate understanding of the amphibians
and reptiles of southern Florida as a fauna, we must examine not
only the distribution of the species in the major habitats, but in the
distinct geographical areas as well. A knowledge of the former helps
to clarify the latter. Essentially southern Florida can be divided
into two geographical areas, the mainland and the Florida Keys.
The geographical distribution of the herpetofauna on the main-
land is a direct reflection of the ecological distribution of the com-
ponent species. The southern part of the peninsula consists of rela-
tively high land along the coasts supporting xeric and some mesic
habitats, and a low interior basin with wet and mesic habitats. Spe-
cies restricted to the dry environments are found only in the eastern
and western coastal regions; those that are restricted to the wet habi-
tats are found only in the interior region. The geographical distribu-
tion of a given species may be ascertained by examining figure 3
and the locality records for that species in the "Accounts of the spe-
cies." Exclusive of the sea turtles, there are 80 species (87 species
and subspecies) of amphibians and reptiles in southern Florida. With
the exception of 3 species of introduced lizards and 3 native sub-
species, all occur on the mainland (see table 2).
The other principal geographical area, the Florida Keys, by nature
of its geological history and present vegetation, may be subdivided
into the upper keys (Key Largo to Bahia Honda Key inclusive) and
the lower keys (from Bahia Honda to Key West). The upper keys are


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BULLETIN FLORIDA STATE MUSEUM


a coral reef formation supporting luxuriant tropical hammocks. The
lower keys have been formed from Miami oblite, the same formation
making up the eastern rim of the peninsula, and support a dry pine-
palmetto association. Twenty-one species of amphibians and reptiles
are definitely reported from the keys, not including introduced spe-
cies. Those species inhabiting the mainland and the upper and
lower keys are:
Hyla cinerea Eumeces egregius
Hyla squirella Eumeces inexpectatus
Gastrophryne carolinensis Diadophis p. punctatus
Kinosternon bauri Coluber constrictor
Terrapene carolina bauri Opheodrys aestivus
Malaclemys terrapin Drymarchon corals couperi
Crocodylus acutus Elaphe g. guttata
Alligator mississipiensis Natrix sipedon compressicauda
Anolis carolinensis Agkistrodon piscivorus
Cnemidophorus s. sexlineatus Crotalus adamanteus
Lygosoma laterale
Of these, Kinosternon bauri, Malaclemys terrapin, Crocodylus acu-
tus, Alligator mississipiensis, Natrix sipedon compressicauda, and
Agkistrodon piscivorus are either restricted to salt- and brackish-
water habitats or occur in these habitats so frequently that dispersal
in Florida Bay and the keys would be accomplished with little diffi-
culty.
Three species are known from the mainland and the upper keys,
but not from the lower keys; these are Heterodon platyrhinos, Tan-
tilla coronata wagneri, and Micrurus f. fulvius. Both Tantilla and
Micrurus are frequent inhabitants of hammocks, a dominant habitat
on the upper keys; Heterodon is usually found in pine woods or open
sandy areas on the mainland; the latter is a restricted habitat on the
upper keys. Six species occur on the mainland and the lower keys,
but not on the upper keys; these include: Scaphiopus holbrooki, Bufo
quercicus, Bufo t. terrestris, Rana pipiens, Storeria dekayi victa, and
Thamnophis sauritus sackeni. With the exception of Rana pipiens,
all of the above species are commonly found in pine woods on the
mainland; this habitat is widespread on the lower keys.
In some species different races inhabit the keys and the mainland.
Eumeces e. egregius occurs on the lower keys and the upper keys;
E. e. onocrepis is found on the mainland. Coluber constrictor priapus
inhabits the lower keys and the mainland north of the Everglades,
whereas the subspecies paludicola lives in the Everglades and the
upper keys. Kinosternon b. bauri is found on the lower keys, the
subspecies palmarum on the upper keys and mainland. Malaclemys


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 215

terrapin rhizophorarum may be restricted to the keys; othre subspecies
occur on the mainland. In most species there are no striking differ-
ences between the populations on the mainland and on the keys;
however, in some, interesting variation has been noted. The popula-
tion of Bufo terrestris on Big Pine Key consists of toads that are
noticeably smaller than ones from the mainland; the individuals from
the keys have a greenish color compared to the brown of the main-
land toads. Continuous clinal variation from northern Florida onto
the keys is apparent in Gastrophryne carolinensis, Elaphe guttata,
and Micrurus f. fulvius. In five species occurring on the lower keys
the range of variation in certain characters approximates more closely
that found in samples from northern Florida than that from samples
from the southern part of the peninsula. In Lygosoma laterale and
Eumeces egregius the populations in northern Florida and on the
keys resemble one another in color pattern. This is also true in Colu-
ber constrictor. Furthermore, in the last species and in Diadophis
punctatus and Storeria dekayi victa, the same pattern of geographical
variation is displayed in certain aspects of the scutellation. Possible
explanations of these patterns of variation and the present geographi-
cal distributions are offered in the following section.

ORIGIN OF THE HERPETOFAUNA 5
With a knowledge of the present geographical and ecological
distributions of the species of amphibians and reptiles in southern
Florida, together with the data presented on the trends in variation,
certain speculations concerning the origin of the herpetofauna may
be made. The questions that may be raised are: What are the
relationships of the fauna? By what dispersal routes did the species
enter southern Florida and the Florida Keys? How were the present
patterns of distribution formed? And, what has been the cause of
differentiation in certain members of the fauna? Needless to say,
none of these questions can be definitely answered. In some instances
where sufficient evidence is available, more than one logical explana-
tion is usable; in others, the picture is so complex and the evidence so
scanty, that little in the way of an explanation can be offered. We
should like to emphasize that, although in most instances we have

While this paper was in press Coleman J. Goin (1958, Comments upon the
origin of the herpetofauna of Florida. Quar. Jour. Florida Acad. Sci. vol. 21,
no. 1, pp. 61-70) presented a discussion of the origin of the present Floridian
herpetofauna on an historical basis. The conclusions he draws are in accord
with those presented here.


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BULLETIN FLORIDA STATE MUSEUM


Vol. 3


substantial bases for the following opinions, these ideas are, for the
most part, purely speculative.
Of the 68 native species of amphibians and reptiles known to
occur in southern Florida, 59 are more or less widely distributed in
the Atlantic coastal plain of the southeastern United States; of these,
many have broad geographical ranges in eastern North America.
Eight species (Pseudobranchus striatus, Kinosternon bauri, Pseudemys
nelsoni, Sceloporus woodi, Eumeces egregius, Neoseps reynoldsi, Sem-
inatrix pygaea, and Liodytes alleni) are restricted to, or center in,
peninsular Florida. Crocodylus acutus occurs in Middle America and
the West Indies, as well as southern Florida. Of the introduced forms,
all but Bufo marinus and Ameiva ameiva (both recent introductions
through human agencies) are widely distributed in the West Indies.
Those introduced species that have more or less broad ranges or that
have undergone differentiation in southern Florida (Eleutherodacty-
lus ricordi, Hyla septentrionalis, Sphaerodactylus notatus, and Anolis
sagrei) may have reached southern Florida from Cuba or the Ba-
hamas by rafting or some other natural means. The other introduced
species have their ranges in southern Florida restricted to port cities
and evidently are recent introductions by man. There is no biologi-
cal or geological evidence that a land bridge ever existed between
the Florida peninsula or the keys and any of the islands in the West
Indies. Thus, with the exception of the introduced species and pos-
sibly Crocodylus, the herpetofauna of southern Florida has been de-
rived directly from that of North America, more specifically from that
of the southeastern Atlantic coastal plain. Of course, many species
inhabiting the southeastern Atlantic coastal plain were derived from
or are related to species or groups of species living in the southwest-
ern United States or Middle America. Thus, Micrurus fulvius, Dry-
marchon corais, and Crocodylus acutus are examples of species hav-
ing wide ranges in Middle America. Others belong to genera which
apparently underwent their major differentiation in Mexico or the
southwestern United States and represent forms that entered the
southeastern Atlantic coastal plain and subsequently peninsular Flor-
ida. Included in this group are such species as Rhadinaea flavilata,
Crotalus adamanteus, Lygosoma laterale, and Cnemidophorus sex-
lineatus. These entered the southeastern Atlantic coastal plain from
the west along the Gulf coastal plain. Evidence for this highway of
dispersal is discussed by Neill (1957) and Auffenberg (1958).
Six genera are endemic or nearly endemic to the Florida pen-
insula, namely: Pseudobranchus, Rhineura, Neoseps, Stilosoma, Semi-


El


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 217

natrix, and Liodytes. Of these Pseudobranchus and Seminatrix have
invaded the Atlantic coastal plain as far north as South Carolina, and
Liodytes as far north as southern Georgia. Of the endemic forms
only Stilosoma and Rhineura do not occur in the southern part of the
peninsula.
To understand the origin and present distributions of these species,
some knowledge of the historical geology of the peninsula is essential.
Most geologists agree that land has persisted in central Florida in the
form of a peninsula or group of islands since Pliocene time and that
with the fluctuation in sea level during the Pleistocene this land
was alternately a peninsula (sometimes more extensive than the pres-
ent one) and an archipelago. Hubbell (1954) has presented paleo-
geographic maps of the Florida peninsula for the Pleistocene. How
much land was emergent in southern Florida, if any, is not known.
There is biological evidence in support of the theory that the east
coast ridge, the lower keys, and parts of what is now Florida Bay
were above sea level, at least as a chain of islands, during the last
part of the Pleistocene. Since the region that is now central Florida
was alternately connected with and separated from continental North
America during the Pleistocene and probably during the Pliocene,
the Florida endemics could have been isolated in that area and differ-
entiated from their parent stocks, or, some of the endemics may be
relict populations of once widespread species that are now extinct
elsewhere. Among those that may have arisen through isolation in
this manner are: Pseudobranchus from an early sirenid stock, Kinoster-
non bauri from an early subrubrum stock, Pseudemys nelsoni from
an early rubriventris stock, and Sceloporus woodi from an early un-
dulatus stock. The other endemic species have no close allies living
in the southern or eastern United States; these relict forms include
Rhineura floridana, Eumeces egregius, Neoseps reynoldsi, Stilosoma
extenuatum, Seminatrix pygaea, and Liodytes alleni. After the con-
nection of the islands with the continent, most of these species re-
mained in the area of central Florida; some followed sandy habitats
or marshes southward, and Pseudobranchus, Seminatrix, Eumeces and
Liodytes moved northward to invade the Atlantic coastal plain.
Until late in the Pleistocene, most or possibly all of southern
Florida and the Florida Keys was submerged. With the emergence
of this land came the development of soils followed by plant and
animal immigrants from the north. With the spread of Pinus caribaea
and Serenoa repens came members of the pineland herpetofauna.
Probably the area that is now the Everglades was part of a large body
of water, of which only Lake Okeechobee remains. With the deposi-


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BULLETIN FLORIDA STATE MUSEUM


tion of muck soils filling in the shallow basin, the prairie flora de-
veloped; then from the north immigrated the members of the prairie
herpetofauna. This seems to be a simple and logical manner of popu-
lating the newly emerged southern part of the peninsula with animals
from the older land mass to the north. It is in regard to populating
the Florida Keys that difficulties arise. The upper keys have an im-
poverished fauna derived from the southern mainland; all of the spe-
cies could have dispersed through the chain of islands by rafting
or other fortuitous means. Likewise, the depauperate fauna of the
lower keys was derived from the mainland. However, 6 native species
occur on the lower keys and on the mainland, but not on the upper
keys. There are certain ecological differences between the upper
and lower keys. First, the upper keys have a predominantly mesic
vegetation consisting of hammocks; the vegetation of the lower keys
is a pine-palmetto association, the same that is present over large
areas of the southern mainland. Second, the lower keys and the
eastern rim of the peninsula are made up of the same geological
formation, the Miami oblite, whereas the upper keys are the remains
of a coral reef built up on the seaward side of the Miami oolite.
Third, the edaphic conditions, apparently controlled by the under-
lying rock, are essentially the same on the lower keys and the eastern
rim of the mainland, but different on the upper keys. Therefore, it
appears that because of the differences in geology, different soils and
consequently different vegetations have developed on the upper and
lower keys.
In southern Florida the Pinus caribaea-Serenoa repens associa-
tion which is widespread in the southern part of the peninsula and
on the lower keys apparently is a fire disclimax. In areas where fires
have been controlled for a number of years mesic hammock has in-
vaded the pine forests. Climatic succession possibly has permitted
mesic hammock to replace pine forest on the upper keys, but not
on the lower keys. Although the soils of the lower keys and eastern
mainland are the same, the temperatures of the lower keys are higher
and the rainfall less than on the mainland. This may account for
the absence of mesic hammocks on the lower keys.
In attempting to solve the biogeographic relationships of the
lower keys the evidence provided by investigations of the historical
geology of the southern part of the mainland and the keys must be
taken into consideration. As stated earlier there is biological and
geological support for the contention that there was a connection
between the lower keys and the eastern mainland that did not in-
volve the upper keys. Storm (1945) and Price (1954) have provided


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 219

geological evidence, and Neill (1957) and Auffenberg (1958) biological
evidence for a connection between the lower keys and the western
mainland across Florida Bay. The Gulf coast of Florida, including
the area that is now Florida Bay is a submergent coast. Geological
evidence points to the fact that there has been an east-west tilt in
peninsular Florida, resulting in a sinking of the western shoreline.
Florida Bay is a broad shallow expanse of water separating the lower
keys, and the upper keys as well, from the southwestern mainland.
Only a slight amount of sinking is required to submerge the entire
area.
On the basis of our knowledge of the present and past geography
of the peninsula and the keys and of the environments of the areas
under consideration the following possible explanations of the differ-
ences in the herpetofauna of the upper and lower keys may be
offered:
1. Fortuitous rafting has brought more species from the main-
land to the lower keys than to the upper keys.
2. Climatic succession has caused mesic hammocks to replace
pine forests on the upper keys, but not on the lower keys,
resulting in the extinction of certain species on the upper keys,
but permitting their survival on the lower keys.
3. Before the upper keys (the coral reef) emerged from the sea,
there was a land connection across Florida Bay from the main-
land to the lower keys, thus permitting continuous animal pop-
ulations throughout the area. Later the area that is now Flor-
ida Bay submerged, isolating populations on the lower keys.
Finally the coral reef emerged to form the upper keys; this
chain of islands was populated by a secondary invasion from
the mainland.
4. Before the upper keys emerged, the east coast rim was emergent
and continuous from the mainland through the lower keys.
There were continuous animal populations throughout the
area. Later the region of the present upper keys submerged
(possibly correlated with the sinking of the western shoreline),
isolating populations on the lower keys. Finally the coral
reef emerged to form the upper keys, which were populated
by a secondary invasion from the mainland.
Several salient features must be discussed with regard to the
above possibilities. Although there is a relatively short waterway
from the southern tip of the peninsula (Cape Sable region) to the
lower keys, it is unlikely that any of the species occurring on the


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lower keys reached those islands by rafting from the tip of he pe-
ninsula. The southern part of the peninsula jutting into Florida Bay
supports one of the largest mangrove swamps in the world. The
species living on the lower keys, but not on the upper ones, are in-
habitants of dry pine-palmetto environments and do not occur in the
mangrove swamps. Rafting from north to south along the Atlantic
coast of the peninsula to the lower keys is not likely, because the
Gulf Stream flows northward in this area.
If climatic succession of mesic hammock over pine forest has
occurred on the upper keys, species of reptiles and amphibians in-
habiting pine forest but not mesic hammock would have been elim-
inated from the upper keys. This would result in isolated populations
of certain species on the lower keys; however, it does not explain
why there are differences between populations on the upper and
lower keys and often similarities between those on the mainland and
the lower keys. Nevertheless, if such succession did occur (and there
is no evidence to the contrary), this may account for some of the
differences between the herpetofauna of the upper and lower keys.
As stated previously, several species inhabiting the lower keys
have coloration, proportions, or scutellation more like populations
of those species in northern Florida than like those in southern Florida
or on the upper keys, if they occur at all on the latter. Such popu-
lation differences occur in enough species so as to make this factor
just as important as the presence and absence of species in deter-
mining the method by which the lower keys were populated. To
explain these differences we must turn to the geological history of
the area. The major clue provided by the geological evidence is that
sometime in the late Pleistocene, probably in pre-Pamlico time (Wis-
consin) a continuous land mass extended from the region of east-
central Florida to the southwestern tip of the Florida Keys. Further-
more, this probably was continuous across Florida Bay to the Gulf
coast of the peninsula, which at that time was west of the present
shoreline. With the rise in sea level this land mass was fragmented
into an archipelago, and the southern part of the peninsula, with
the possible exception of the eastern rim, and Florida Bay were
submerged together with the area that is now the upper keys. This
resulted in the complete isolation of the lower keys from the main-
land. In all probability the eastern rim and lower keys composed
of Miami oolite were higher with respect to present sea level than
they are now. This limestone formation has formed a typical karst
topography, showing all of the signs of considerable solution and
sinking, resulting in continuous lowering of the land to its present


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 221

level. In post-Wisconsin time the coral reef rose to form the upper
keys and to provide an insular connection between the mainland and
the lower keys.
How did these geological events affect the distribution and differ-
entiation of members of the herpetofauna? In pre-Pamlico time
continuous populations extended from northern Florida southward
throughout the area of the Florida Keys. With the rise in sea level
and submergence of large areas of land, populations of many species
were isolated in the area that is now the lower keys. Here some
species differentiated slightly from the parental stocks to the north
on the mainland; others apparently remained unchanged. With the
establishment of an insular connection between the mainland and
the lower keys, species moved southward from the peninsula and
onto the upper keys; some continued onto the lower keys where they
encountered members of previously isolated populations. Clinal vari-
ation in many species shows continuous gradual differences from
north to south in the peninsula and often onto the upper keys followed
by an abrupt change on the lower keys, the animals on the lower
keys being more like those in the northern part of the peninsula than
in the southern part or on the upper keys. Some species did not im-
migrate to the upper keys; some of these were species that had
ranged southward to the area of the lower keys in pre-Pamlico time.
This has resulted in the isolated populations of such species as Scaphi-
opus holbrooki, Bufo quercicus, Bufo terrestris, and Storeria dekayi
on the lower keys.
We feel that it is safe to assume that the areas comprising the
eastern rim, the keys, and the western flatlands were emergent and
supporting vegetation and most of the species of the present herpeto-
fauna before the southern part of the Okeechobee basin filled with
sediments and formed the Everglades and associated prairies and
savannas. The herpetofauna of the Everglades is similar to that of
the drier Kissimmee prairies to the north of Lake Okeechobee and
probably was derived from the fauna of that area. Some of the most
striking differentiation in the herpetofauna of southern Florida cen-
ters in the Everglades. Here are found the apparently ecogenotypic
populations of blue-gray Coluber constrictor, the orange-colored
Elaphe obsoleta, the pale Lampropeltis getulus, and the pale Kinoster-
non bauri, together with other species showing differences in size
and proportions. Most of these species are inhabitants of the eastern
rim as well as the Everglades. Usually populations on the rim north
and east of the Everglades and on the lower keys are similar to one


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BULLETIN FLORIDA STATE MUSEUM


another, but different from those inhabiting the Everglades. On the
southern part of the rim and on the upper keys are found populations
with intermediate characteristics. Because of the lack of barriers
to most species, there is gene flow between populations in the Ever-
glades and the older, undifferentiated populations on the rim. This
results in populations from the Everglades interdigitating with those
from the rim, populations showing intermediate characteristics, or a
complex mosaic of populations having recombinations of character-
istics or even new characteristics.
Because the Everglades serve as a barrier to many inhabitants of
the pine forest and scrub habitats, there is some isolation of these
populations in southern Florida. Populations of Hyla femoralis, Hyla
gratiosa, Sceloporus woodi, and Gopherus polyphemus in the scrub
habitats near Naples, although not geographically remote, are widely
separated in terms of gene flow from those populations in northeastern
Dade County; the scrub habitats are absent from the interior of the
peninsula south of Lake Okeechobee. Some minor differences are
found between the populations to the west and those to the east of
the Everglades. These are discussed specifically in the accounts of
the individual species.

ACCOUNTS OF THE SPECIES
In the following pages the 92 forms of amphibians and reptiles
known to occur in southern Florida are discussed in relation to their
variation and differentiation in that area, their life histories, and their
ecological and geographical distributions in the southern part of the
Florida peninsula and the keys. The accounts of geographical dis-
tribution are based on 8755 museum specimens from southern Florida.
Of these, data pertaining to variation and differentiation are pre-
sented for 3116 specimens. Comparative material examined from
northern Florida and from other parts of the Atlantic coastal plain,
and from various Caribbean islands consists of 2657 specimens.
Locality records for each form are given at the end of the account.
Mileages are road distances, and directions are road directions, un-
less otherwise indicated. Thus, the Tamiami Trail extends due west
from Miami; 19 miles west of Miami a road leaves the Tamiami
Trail and extends south. A specimen collected 5 miles south of this
road junction is listed as: Dade County: 19 mi. W, 5 mi. S of Miami.
Museum catalogue numbers have been used only when reference
is made to a particular specimen in the text; the abbreviations used
for the museums are:


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 223

AMNH American Museum of Natural History
ANSP Academy of Natural Sciences of Philadelphia
CAS Chicago Academy of Sciences
ChM Charleston Museum
CM Carnegie Museum
CNHM Chicago Natural History Museum
CU Cornell University
DRP Dennis R. Paulson, Miami, Florida
ERA-WTN E. Ross Allen-Wilfred T. Neill, Silver Springs, Florida
MCZ Museum of Comparative Zoology
SU Natural History Museum, Stanford University
UF University of Florida Collections
UI University of Illinois Museum of Natural History
UMMZ University of Michigan Museum of Zoology
UMRC University of Miami Reference Collection
USNM United States National Museum
Means of counts and measurements are in parentheses after the
observed range, for example, 91-112 (104). Capitalized color names
are from Ridgeway (1912). Unless otherwise indicated by the pres-
ence of a synonymy, all names are in accordance with the sixth edition
of the "Check list of North American amphibians and reptiles" by
Schmidt (1953). Literature references in the synonymies which are
not complete are contained in the "Literature cited."

Siren lacertina Linnaeus
In life the coloration is grayish above; the ventral surface is irreg-
ularly mottled or spotted with pale green. These colors fade rapidly
in preserved material, and the color shortly becomes a dull uniform
gray. The largest individual from southern Florida, a female, has a
snout-vent length of 452 mm. and a total length of 611 mm. Most
specimens are of a much smaller size, and very large adults rarely
have been taken in southern Florida. The ratio of total length to
tail length in 38 specimens varies from 2.6 to 3.8 (3.0). The number
of costal grooves varies from 33 to 40 (37.0). Data from seven speci-
mens from northern Florida and 31 from South Carolina show that
in those from South Carolina the ratio varies from 3.0 to 3.7 (3.2),
and the number of costal grooves ranges from 36 to 39 (37.4). In the
specimens from northern Florida the total length to tail ratio varies
from 2.7 to 4.2 (3.0), and the number of costal grooves varies from
38 to 40 (38.6). Goin (1942: 217) pointed out that the number of
costal grooves is of primary importance in distinguishing S. lacertina
from S. intermedia. He stated that S. lacertina has on the average 38
costal grooves, while intermedia varies from 31 to 38. The data pre-
sented here show that populations of S. lacertina have costal grooves




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224 BULLETIN FLORIDA STATE MUSEUM Vol. 3

varying in number from 33 to 40, a range generally overlapping that
of intermedia; however, the coloration of the two species allows for
easy identification of fresh specimens. S. intermedia has not been
taken in southern Floirda.
Neill (1949a) described and differentiated between the juveniles
of S. lacertina and intermedia, basing his conclusions on specimens
collected near Midville, Burke County, Georgia. The young of both
species were collected together in a shallow flatwood pond; Neill
stated that S. intermedia normally breeds and lives in such ponds,
and that the young of S. lacertina probably had been washed into the
pond by a freshet from a nearby river. Neill stated that juvenile
S. intermedia possess a bright red band across the snout that extends
irregularly almost to the gills; the rest of the animal is grayish-brown
above and below. The illustration of this juvenile shows the tail fin
pigmented (except for a narrow band on the tail itself) and extending
anteriorly on the body for about one quarter of the snout-vent length.
On the other hand, juveniles of S. lacertina are characterized by a
light yellowish stripe on the side from the base of the gills to about
the level of the vent; a less distinct yellowish ventrolateral line also
is present. The caudal fin, which extends far anteriorly on the body,
is described as clear yellowish white. There is an irregular brownish
line from the eye to the base of the gills.
Comparison of 20 juveniles from southern Florida with the above
descriptions indicates that they resemble Neill's S. lacertina in some
ways, but disagree in others. The tail fin is heavily pigmented with
dark brown; it is not clear and does not extend anteriorly as described,
but terminates immediately anterior to the vent. There are no lat-
eral or ventrolateral yellowish stripes as described by Neill, nor were
such stripes present in the living specimens. Usually there is a faint
postocular stripe, and the labial area is pale brown. The top of the
head has a few scattered dark spots on a gray ground color. These
juveniles vary in snout-vent length from 33 to 49 mm. The possi-
bility that these juveniles represent S. intermedia has not been over-
looked. Their designation as S. lacertina is based on the fact that
they lack (now and when collected) any red markings as described
by Neill for S. intermedia, and that they were taken at the same lo-
cality where adult S. lacertina were secured. It would seem that if
these juveniles represent S. intermedia, the adults of the species also
would have been encountered. Such is not the case. It is possible
that juvenile sirens may vary in details of color pattern at different
localities, and thus result in the discrepencies noted.


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 225

The majority of specimens of S. lacertina were secured by net-
ting and sieving water hyacinths in the Tamiami Canal at a period
of low water. The adults were taken in deep water (21/ to 3 feet),
whereas juveniles occurred principally in a shallow (6 to 12 inches)
mucky-bottomed pond. One was observed in water beneath a piece
of limestone. During periods of very high water, for example 1947,
the Tamiami Trail was locally flooded and sirens were observed to
cross the road. Individuals were taken from tunnels in canal banks.
These tunnels, which slant downward at about 45 degrees and then
level off, were 14 to 19 inches long and 1/4 to 21/ inches in diameter.
At the end of each tunnel was a chamber about 6 inches in diameter
in which the animal rested and where it was usually captured. The
tunnel openings were about a foot below the water surface. During
periods of high water Siren are difficult to secure since at that time
the salamanders occupy whatever water is available and are not
restricted to sloughs and canals as in periods of dryness. That sirens
occupy temporary water is shown by the collection of specimens
in a temporary shallow pond in Broward County. They are pre-
sumed to gain access to such ponds by tunneling through the mucky
substrate.
The stomachs of 9 specimens have been examined to determine
food habits. These stomachs contained the following: 1 crane-fly
(Tipulidae), 6 rat-tailed maggots (Stratiomyidae), 4 water beetles
(Hydrophilidae), 2 scarabid beetles, 4 nymphs (Odonata: Aeschnidae
and Libellulidae), 1 mayfly nymph (Ephemeridae), 2 crayfish (Astaci-
dae), and 1 fish (Gambusia holbrooki). This last food item indicates
that at least occasionally Siren lacertina does eat fish, contrary to the
opinion expressed by Goin (1957: 41).
The Everglades is the principal habitat for sirens, and the species
has been taken frequently from the canals in this region. No speci-
mens are at hand from cypress ponds, although it is likely that the
siren occurs there. This salamander is absent from the keys. Lo-
cality records for 97 specimens are:
Dade County: Everglades near Miami (1), 20.1 mi. W Miami (29), 20.6 mi.
W Miami (1), 21 mi. W Miami (22), 23.1 mi. W Miami (42), Tamiami Trail bridge
no. 42 (1). Monroe County: Pinecrest (1).

Pseudobranchus striatus belli Schwartz
Pseudobranchus striatus belli Schwartz, 1952a, p. 1.
Eighteen large individuals vary in total length from 107 to 153
mm. There are 29 to 33 (31.5) costal grooves. A detailed description


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BULLETIN FLORIDA STATE MUSEUM


of coloration and a comparison with other forms is to be found in
Schwartz (1952a).
The only individuals of this species collected in the area under
consideration were taken from side canals south of the Tamiami Trail
during times when these canals were temporarily cut off from the main
canal and when the water in them was relatively low. The majority
of specimens was secured by sieving decaying masses of water hya-
cinths and examining the debris. What the "normal" habitat of
these salamanders is can be only conjectured. Neill (1951b) com-
mented on the habitat preferred by P. s. lustricolus in the Gulf Ham-
mock region. Here, where the water hyacinth has not been intro-
duced, the salamanders inhabit the muddy bottoms of pools. Pre-
sumably Pseudobranchus occupy a similar niche in southern Florida,
but they may be more readily secured where they occur in dead
hyacinths.
The food of Pseudobranchus s. belli consists of aquatic oligochaetes.
Of 12 topotypes examined, 4 contained worms of the family Lum-
briculidae, while 8 others contained worms of the family Megascoleci-
dae (Sparganophilus). The type of food eaten indicates that foraging
takes place among the roots of the water hyacinths.
This species is known only from a few localities in Dade County.
It is presumed to occur throughout the Everglades; however, its
aquatic habits and the general inaccessibility of the area make speci-
mens difficult to secure. In northern and central Florida other forms
of Pseudobranchus striatus have been collected in a variety of aquatic
habitats, even in isolated cypress ponds. It should be looked for in
such situations near Naples, Collier County. Flooding undoubtedly
aids in the dispersal of the species. Locality records for 83 speci-
mens are:
Dade County: Lemon City (1), 20.1 mi. W Miami (1), 23.1 mi. W Miami (81).
Notophthalmus viridescens piaropicola Schwartz and Duellman
Diemictylus viridescens piaropicola Schwartz and Duellman, 1952, p. 219.
Diemictylus viridescens evergladensis Peterson, 1952, Herpetologica, vol. 8, pt. 3,
p. 103.
Notophthalmus viridescens piaropicola, Smith, 1953, Herpetologica, vol. 9, pt. 2,
p. 98.
Adults range in total length from 79.0 to 104.0 mm. and from
37.2 to 55.9 mm. in snout-vent length; females average slightly larger
(91.5 mm. total length, 43.5 mm. snout-vent length) than do males
(88.3 mm. total length, 41.4 mm. snout-vent length). The dorsum


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 227

is dark (Fuscous-Black) in life and after preservation, and the Buff-
Yellow belly, strongly mottled with black, is noteworthy. This sub-
species, which ranges throughout the Florida peninsula from Citrus
County southward, is compared with others in Schwartz and Duell-
man (1952).
The type series of N. v. piaropicola was taken by dipping the
submerged masses of Utricularia foliosa, Naiis flexuosa, and Hydro-
trida caroliniana from a drying cypress pool. Large numbers of these
newts also were secured by dipping water hyacinths-both dead and
alive-from canals; specimens also were obtained in isolated ponds,
away from the main body of the Everglades. Schwartz and Duell-
man (p. 226) mention the taking of specimens beneath the lodges of
Neofiber alleni; occasional specimens have been taken from beneath
partially imbedded pieces of oilitic limestone along the Tamiami
Trail.
At the time of the description of N. v. piaropicola, no efts from
southern Florida were known. Examination of material in the United
State National Museum indicates that a specimen there (85329) is
an eft. It was taken by M. K. Brady at Pinecrest, Monroe County,
14 February 1932; it has a total length of 45 mm. and a snout-vent
length of 21 mm. The skin of the dorsum is highly granular, light
brown in color, and is spotted with brown like the venter, although
less so. The tail is terete, and gills are absent. Neotenic individuals
have been pointed out by Schwartz and Duellman (p. 225); these are
similar in color and pattern to the adults.
Females taken in the spring and early summer may have eggs,
and this indicates that egg deposition probably occurs at least during
this period. However, it is likely that there is a more extensive breed-
ing season. Small larvae (USNM 85330-85346) were taken 21 January
1932, in potholes which contained Isnardia; this winter date tends to
confirm a lengthy reproductive season.
The stomach contents of 19 topotypes were examined. The most
abundant material was newt skin (13 individuals, 59% of bulk).
Decapod crustaceans (cf. Atylidae) occurred in 8 stomachs and
formed 31.3 percent of the bulk. Amphipods, Diptera (Tanypodinae,
Chironominae), Odonata (Anisoptera), and Gastropoda (Gyraulus)
also were eaten. Thirteen specimens from another locality also were
examined; all contained algaelike material, and two specimens con-
tained fragwt. of insects (Coleoptera: Hydrophilidae and Dytisci-
dae).
Notoplitlialinis has been taken from canals and sloughs along the
Tamiami Trail and in the Everglades. Individuals may be abundant


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in cypress ponds and willow-bordered ponds at the margins of the
Everglades. The species is presumed to occur throughout southern
Florida with the possible exception of both eastern and western
pinelands, where it may occur locally in mesic situations. Locality
records for 905 specimens are:
Collier County: 2.3 mi. W Collier-Dade Co. line on Tamiami Trail (5), 5.2
mi. E Monroe Station (81). Dade County: 3.5 mi. SE Collier-Dade Co. line on
Tamiami Trail (7), Loop Road (2), 19 mi. W Miami (5), 20.1 mi. W Miami (413),
20.6 mi. W Miami (26), 21.2 mi. W Miami (47), 22 mi. W Miami (38), 23.1 mi. W
Miami (176), 24.9 mi. W Miami (5), 26 mi. W Miami (53), Paradise Key (13), 1.3
mi. N, 5.2 mi. NE of Paradise Key (6). Monroe County: Flamingo (1), 60 mi. W
Miami (5), 5.4 mi. S Monroe Station (20), Pinecrest (2).

Amphiuma means Garden
Amphiuma means Garden, 1821, in Smith, Correspondence of Linnaeus. vol. 1,
pp. 333, 599. Hill, 1954, Tulane Studies Biol., vol. 1, no. 12, pp. 191-215.
Amphiuma means means, Goin, 1938, Herpetologica, vol. 5, pp. 127-130.
In 49 specimens from Dade County the number of costal grooves
varies from 51 to 61 (57.6); the vomerine teeth vary from 25 to 46
(33.3). In total length the largest individual measured 625 mm., and
the smallest 93 mm.; the three largest males measured 385, 399, and
404 mm., and the three largest females measured 391, 470, and 625
mm. The color of both young and adult individuals after preserva-
tion is black dorsally, grading to Deep Neutral Gray ventrally. Nei-
ther in life nor after preservation can these specimens be called
Mummy Brown, as stated by Bishop (1943: 53). The belly color of
some immature specimens is lighter (Deep Mouse Gray). The dor-
sal and ventral colors gradually merge into each other and there
is no clear line of demarcation between them. The upper and lower
jaws are black, and the fleshy flanges on the lower jaw are grayish.
The ratio of total length to body length in 55 southern Florida
specimens varies from 1.2 to 1.4 (1.32). This may be compared to
the ratio in six specimens from South Carolina in which the range of
variation is from 1.2 to 1.3 (1.27); three specimens from Alachua
County, Florida have a mean ratio of 1.30. The ratio of length of
tail to total length in 55 specimens from southern Florida varies
from 19.7 to 28.2 (24.7), in six specimens from South Carolina from
19.3 to 23.8 (21.6), and in three specimens from Alachua County,
Florida from 23.6 to 26.4 (25.1). Specimens from South Carolina
have 31 to 44 (38.4) vomerine teeth, a somewhat higher average num-
ber than that found in southern Florida. Obviously, additional ma-


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 229

trial is needed from the northern part of the range before adequate
assessment of the data can be made. The impression is, however,
that at least the length of the tail varies clinally, the individuals with
shorter tails being in the northern part of the range.
Amphiuma is strictly aquatic and consequently is restricted to
permanent standing water, or water with a low flow gradient. As far
as known, this salamander does not occur in rivers in southern Flor-
ida, and it is absent from water which may be under saline influence
during yearly fluctuation. Most southern Florida specimens have
been secured by removing water hyacinths from canals and sloughs
and examining the roots thereof. In this manner Amphiuma has
been taken with Pseudobranchus, Siren, and Notophthalmus. How-
ever, at the type locality of the last, a cypress pond, no Amphiuma
were secured.
These salamanders form a major part of the diet of Farancia
abacura; on two occasions this large snake has been seen in a violent
struggle with a large salamander, presumably Amphiuma, but pos-
sibly Siren.
Examination of eleven stomachs of Amphiuma yielded the follow-
ing results: Two were empty; three contained mucus and plant frag-
ments. The other six stomachs contained 4 rat-tailed maggots (Syrphi-
dae), 2 naiads (Libellulidae), 5 aquatic beetles (Dytiscidae and Hydro-
philidae). Two individuals had eaten crayfish, and one stomach
contained only the remains of a fish (Mollienesia latipinna). A spe-
cimen kept in captivity was fed Fundulus, Chryopeops, and Jordan-
ella, and was seen to eat the eggs directly from the swimmerets of a
crayfish, and later to eat the crayfish itself. Hamilton (1950) ex-
amined the stomachs of Amphiuma from Ft. Myers, Florida, and
North Charleston, South Carolina; he found that in a combined sample
of 36 specimens the food taken included insects (52.8% in frequency
and 33.8% in bulk) as the most important item, and that the sala-
manders also ate amphibians, reptiles, fish, crustaceans, mollusks, and
arachnids.
Weber (1944) reported on a female Amphiuma and her eggs. The
nest was discovered on 3 February 1933 at a locality 25 miles south
of Royal Palm Hammock (now Paradise Key), Dade County. The
female (390 mm. in length) was found under a board imbedded in
muck, and was coiled around 49 eggs. The eggs were in a rosary-
like string. A second set of eggs was received by Weber on 10 Feb-
ruary 1935, and these were hatched on 21 June 1935. Assuming
that the eggs are deposited in mid-January, the incubation period









BULLETIN FLORIDA STATE MUSEUM


is estimated to be about five months. It is presumed that the find-
ing of a female Amphiuma with her eggs represents a fortuitous oc-
currence in which the object under which the female burrows to lay
the eggs is exposed by fluctuations in water level. Since the respira-
tion in these forms is primarily pulmonary, such exposed nesting sites
are able to remain inhabited as long as they remain moist and do not
become overheated. In this regard, Weber stated that the board
under which the brooding female was taken "was covered with the
dry and caked marsh scum and decayed vegetation. This covering
apparently acted as an insulator against the baking action of the sun
and conserved moisture for the eggs." Baker (1945: 64), in discussing
these egg-laying activities, stated that "burrowing into the mud
bank" would better express the terrestrial activities of Amphiuma
than "crawling onto land."
Amphiuma probably is distributed over the central part of south-
ern Florida, in the Everglades region. There are no records from the
east coast ridge (where suitable habitat is limited), nor from the
keys, where it certainly does not occur. In the western region it
probably occurs in canals and cypress ponds, but specimens and rec-
ords are lacking. Locality records for 69 specimens are:
Dade County: Coral Gables (1), 4.2 mi. S Florida City (2), Miami (2), 20.1
mi. W Miami (35), 23.1 mi. W Miami (23), Paradise Key (1), 25 mi. SW Paradise
Key (1), Tamiami Trail bridge no. 42 (2), Monroe County: Pinecrest (2).

Scaphiopus holbrooki Harlan
Rana holbrookii Harlan, 1835, Medical Physical Research, p. 105.
Scaphiopus albus Garman, 1877, Proc. Amer. Assoc. Adv. Sci., vol. 25, p. 194.
Scaphiopus holbrooki albus, Carr, 1940, p. 53.
Scaphiopus holbrooki, Duellman, 1955a, p. 143.
Adult males vary in snout-vent length from 45 to 64 mm.; females
are slightly smaller, ranging from 43 to 63 mm. Specimens from
Key West, formerly considered to be a different subspecies, are
smaller than those from the mainland, the males averaging 51.7 mm.
in snout-vent length as compared to 59.2 mm. for the males from
the Miami area. Typical individuals from the mainland have a light
tan or grayish dorsal ground color with three more or less distinct
dark brown or olive-brown longitudinal bands. Many individuals
from the lower keys are lighter in color. The variation in this species
with respect to the Florida populations has been discussed at length
by Duellman (1955a: 141-3). Aside from the slight variation in the
intensity of the dorsal pigmentation, the only observable trend is in


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 231

the over-all size of the individuals, there appearing to be a trend
toward smaller size from north to south in the Atlantic coastal plain,
peninsular Florida, and the Florida Keys.
Scaphiopus occurs in the sandy regions on the eastern rim of
southern Florida, being found in the palmetto scrub and pineland.
Specimens also have been found at Paradise Key, Dade County, a
locality presenting in general more mesic conditions than the eastern
rim. The abundance of the spadefoot may be easily underestimated,
for nothing short of torrential rains will bring them from their under-
ground retreats. For this reason many residents refer to them as the
"hurricane frog." Their abundance in vacant lots, on lawns, and in
the streets of cities after a heavy rain is surprising. Carr (1940: 53)
stated that in Key West the species was most abundant in vacant lots
on the south edge of the island.
Individuals have been found in their burrows, which are nearly
one and one-half inches in diameter and at least eight inches in
length. Spadefoots were observed emerging from such burrows
during a heavy rain in Miami.
On three different occasions when there has been rainfall in
excess of three inches within a twelve-hour period, large breeding
choruses have been encountered. Two of the choruses were in the
northwestern part of Miami on 5 and 27 October 1952. The other
was encountered near Surfside, Dade County, on 27 June 1953. The
following notes concern the chorus of 5 October 1952. Scaphiopus
were seen in an open marshy field that was flooded in many places
due to a very heavy rain. At 2:30 P.M. many individuals were seen,
but none was calling. Within an hour after the first males began
calling, the chorus of an estimated 2000 individuals put forth an ear-
splitting racket. As the rain continued to fall, the spadefoots in-
creased in numbers. In the largest pond, which extended for nearly
a mile and was approximately 300 feet wide, there were at least four
individuals per 100 square feet, giving an estimated total of more than
60,000 spadefoots in this large pond. In the same area Scaphiopus
were calling from every pool of water, including small holes in the
road and even from a flower pot! There were several other large
choruses in the vicinity, and there are reliable reports that the species
was calling in many other places in Miami. The following night only
a few scattered individuals were calling.
The males call from the open water and float with their legs out-
stretched. When the vocal sac is inflated the forepart of the body
is reared upward at an angle of about 75 degrees and the legs are


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232 BULLETIN FLORIDA STATE MUSEUM Vol. 3

forced backward. The call is a nasal "waank" coming about once
every two seconds. These calls are repeated from 6 to 15 times, after
which there is a period of silence for about 20 seconds. Males were
observed to clasp other males, which caused the defendant to spin
in a circular motion and call rapidly until the offender released him.
A male Scaphiopus was found in amplexus with a female Bufo ter-
restris.
Many small females were observed in the congregation, but in-
dividuals measuring less than 43 mm. in snout-vent length were not
gravid. The eggs are laid in an elongated mass that is almost string-
like in appearance; these are attached to sticks and weeds six to
eight inches below the surface of the water. There are no further
data available concerning the breeding habits of this form in south-
ern Florida. The length of time required from hatching to meta-
morphosis is not known, although it probably coincides with that
given by Wright and Wright (1949: 126) who give the minimum time
as 14 days. A series of 63 recently transformed young from Prince-
ton, Dade County, average slightly more than 13 mm. in body length.
Food taken from the stomachs indicates that the diet consists
mainly of terrestrial arthropods, including beetles, ants, grasshoppers,
Dermaptera, and spiders. One terrestrial gastropod was found. One
specimen collected in the chorus described above regurgitated an
Acris gryllus, and another, a Bufo quercicus.
In southern Florida Scaphiopus is found only in the eastern part
of the peninsula and on the lower keys. Wright and Wright (1949:
129) report this species from Matecumbe Key; however, since there
are no specimens from that locality, the record is open to question.
Locality records for 412 specimens are:
Broward County: Dania (1). Dade County: Biscayne Gardens (1), Coral
Gables (1), Little River (1), Miami (204), 12 mi. W, 2.1 mi. S Miami (1), 19 mi.
W, 9.5 mi. S Miami (1), Miami Beach (5), Paradise Key (21), Princeton (63), South
Miami (1), Surfside (23). Monroe County: Key West (89).

Bufo marinus Linnaeus

Rana marina Linnaeus, 1758, Systema Naturae, ed. 10, vol. 1, p. 211.

This species has been introduced into the Miami area and appar-
ently has become established. L. Neil Bell (in litt. to Roger Conant)
stated: "I observed mated pairs, tadpoles, and about 11 calling males
at one little pond. They were calling elsewhere in the general vi-
cinity." Dennis R. Paulson stated (in litt.) that in the summer of 1957


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 233

the population was thriving. One specimen collected by Bell (UMMZ
113000) has the following data: western part of Miami, Dade County,
17 May 1955. The specimen is a male with a snout-vent length of
140 mm.
Several years ago Bufo marinus was introduced into the sugar
cane fields near Pennsuco, which is about 10 miles northwest of
Miami. Possibly the present population was derived from that in-
troduction which supposedly died out. More likely, however, the
present population has resulted from either accidental or intentional
release of captive specimens by animal dealers in the Miami area.
The single locality record is cited above.

Bufo quercicus Holbrook
A series of 20 males from Miami average 26.5 mm. in snout-vent
length; six females average 27.9 mm. From the small number of adult
specimens available from Big Pine Key, it appears as though the indi-
viduals on the keys are somewhat smaller than those on the main-
land; both sexes average 24.2 mm. in snout-vent length on Big Pine
Key. The largest specimen examined, a female, has a snout-vent
length of 30 mm.; the smallest juvenile measured 14 mm. The dorsal
ground color is a light tan. The dorsal markings are dark brown, and
the flanks are mottled with dark brown and cream. The middorsal
light line is golden yellow or orange. The venter usually is clear
cream color, although in some the ventrolateral areas are grayish,
and in a few specimens the belly is mottled with black. The speci-
mens from the keys are generally darker with a thinner and more
dull-colored middorsal line (fig. 20).
Specimens from southern Florida were compared with series
from Silver Springs, Marion County, Florida, and from Charleston,
South Carolina. The Charleston specimens are slightly darker than
the Florida ones, the middorsal line being dull in color and the
venter dusky. The specimens from Silver Springs are essentially the
same as those from southern Florida.
This species is characteristic of the rocky and sandy pineland and
is commonly found in areas of sandy scrub. It also occurs in ham-
mocks and in the wet prairies, but it reaches its greatest abundance
in the pineland. It is often encountered during the day. Individuals
were found to be active at midday in the sandy scrub areas on Marco
Island. Scattered males have been heard calling during the day.
Choruses have been heard from mid-April to late October, and pre-
sumably breeding takes place between those dates. The breeding


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Vol. 3


choruses usually are found in small pools and shallow ditches, the
toads never being found in deep water. The males, more often than
not, call from small clumps of vegetation at the edge of the water,
although amplexus takes place in the water. The call is a shrill
chicklike "peep."


*A 11




Figure 20.-Variation in the dorsal color pattern in Bufo quercicus from
Miami.
Examination of the stomach contents of 15 individuals collected
in Miami revealed that ants were the most abundant food item and
made up about half of the diet. Other food items found include sev-
eral beetles, 3 lepidopteran larvae, 1 hemipteran, and 2 spiders.
Bufo quercicus probably constitutes the greatest part of the diet
of small Heterodon. Thamnophis sauritus also have been seen feeding
on this toad.
In southern Florida Bufo quercicus is abundantly found on the
east coast rim and the Gulf coastal region as far south as Marco
Island. Less frequently it is encountered in the Everglades. It is
also known from Big Pine Key and Boca Chica Key; presumably it
occurs on all of the lower keys. Locality records for 439 specimens
are:
Broward County: 1 mi. W Davie (17), 1.6 mi. W Hollywood (1), 2.2 mi. W
Hollywood (1). Collier County: Marco Island (12), Naples (4), 1.7 mi. SE Naples


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 235

(1), 3 mi. SE Naples (10), 3.2 mi. SE Naples (1), 5 mi. SE Naples (1), 6.7 mi. SE
Naples (28), 7.4 mi. SE Naples (3), 10 mi. SE Naples (1). Dade County: Coral
Gables (2), 1.5 mi. SE Florida City (16), Homestead (2), 4 mi. S Homestead (14),
6 mi. S Homestead (1), Long Pine Key (4), Miami (122), 7 mi. W, 2.5 mi. N Miami
(9), 8 mi. W, 3 mi. N Miami (26), 12 mi. W, 1 mi. N Miami (40), 17.5 mi. W
Miami (15), Miami Beach (1), Miami Springs (11), Opa-locka (16), Paradise Key
(2), Richmond Air Base (44), Uleta (7). Monroe County: Big Pine Key (26), Boca
Chica Key (1).
Bufo terrestris terrestris Bonnaterre
Apparently Bufo terrestris reaches its maximum known size in
southern Florida. The largest male examined had a snout-vent length
of 67 mm.; the largest female, 107 mm. Wright and Wright (1949:
201) gave 92 mm. as the maximum size for this toad. That the above
female is not an exceptional giant is shown by two other specimens
having snout-vent lengths of 104 and 106 mm. The smallest juvenile
is from Big Pine Key; it has a snout-vent length of 9.5 mm. A series
of 11 juveniles from Snake Bight, Monroe County, range in size from
10.0 to 12.7 (10.8) mm. Of the specimens known from the keys, the
largest is a female measuring 70 mm. in snout-vent length. There is
a slight size difference in specimens from the Everglades as compared
with those from the eastern rim; individuals from the lower keys are
notably smaller than those from the mainland (table 3).

TABLE 3
GEOGRAPHICAL VARIATION IN Bufo t. terrestris IN SOUTHERN FLORIDA.
(MEASUREMENTS IN MILLIMETERS.)

Locality Sex N Body Tibia Tibia length Head Head width
length length Body length width Body length

Coral Gables $ 18 58.8 22.5 38.3% 23.3 39.6%
9 13 67.0 23.7 35.4 26.9 40.1
Everglades S 11 58.7 21.0 35.8 22.0 37.5
9 12 65.1 22.0 33.8 24.7 37.9
Big Pine Key $ 5 48.8 14.4 35.7 18.1 37.0
9 24 51.3 18.2 35.5 20.3 39.7

Specimens from the mainland have a light grayish-brown or yel-
lowish-brown dorsal ground color. The dorsal blotches are dark
brown or olive brown, and in a few individuals they are greenish.
In many instances the blotches are large and numerous, with only
thin streaks of the light ground color between them. The venter is
a clear cream color except for the throat of the males, which is black.
Individuals from Big Pine Key had in life, a ground color of light


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BULLETIN FLORIDA STATE MUSEUM


Vol. 3


gray, but dark gray in some and tannish in a few. The dorsal dark
spots were of varying shades of olive green. The parotoid gland was
reddish brown to greenish gray.
Bufo terrestris occurs in the sandy and rocky pinelands, the sandy
scrub, mesophytic hammock, and a variety of edificarian and ruderal
situations. Although found in the wet prairies, it is not nearly so
abundant there as it is in the more xeric habitats.
This species is more nocturnal than Bufo quercicus; however,
during and just after daytime showers it is possible to see these toads
moving about. On three different occasions during the summer of
1951 male Bufo terrestris called from a pond on the University of
Miami campus in Coral Gables following afternoon showers. Early
on a clear morning after a hard rain during the night, this species was
heard calling in a flooded field near Cutler, Dade County. B. terres-
tris breeds in nearly any situation where water is available; they
have been found chorusing in flooded fields, roadside ditches, sink
holes, permanent ponds, canals, sloughs, and even puddles. This
species prefers deeper water than B. quercicus, and where the two
species are breeding in the same body of water, B. terrestris is found
calling and mating toward the middle of the pond, while B. quercicus
stays in the shallow water at the periphery. The breeding season
begins with the first heavy spring rains in April and continues through
August; a few individuals have been heard as late as 27 October.
Their high-pitched trill is often heard on warm summer nights when
there has been no rain.
Recently metamorphosed individuals have been found from mid-
June to early October. On Snake Bight, a narrow spit of sandy marl
surrounded by mangroves and jutting into Florida Bay east of Cape
Sable, scores of juvenile Bufo terrestris were found on the afternoon
of 21 June 1953. The lack of fresh water in the vicinity at that time
poses the question: What is the tolerance of Bufo eggs and tadpoles
to the brackish water in the mangrove swamps?
Several large adults were collected at night under street lights in
the town of Everglades, Collier County. The toads were feeding
on the insects that hit the lights and fell to the ground. At the base
of each lamp post was a six- by eight-inch opening. Several toads
were seen entering and emerging from these holes, a probable day-
time retreat for these city dwellers. An examination of the stomach
contents of these toads revealed that about 95 percent of the bulk
of the food consisted of beetles of the families Carabidae, Cicindeli-
dae, and Curculionidae. Other food items included one grasshopper


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 237

(Tetrigidae), one wasp (Sphecidae), several ants (Formicidae), two
Hemiptera, one Dermaptera, and one small gastropod.
Food items found in the stomachs of ten specimens from Miami
include many beetles (Carabidae, Curculionidae, Elateridae, and Scar-
abaeidae), several ants (Formicidae), two Dermaptera, and one iso-
pod. The beetles accounted for more than 90 percent of the bulk,
and curculionids alone made up about 50 percent of the bulk.
With the exception of the upper keys, Bufo terrestris is found
throughout southern Florida. It is most abundant on the eastern
rim, but is found commonly in the Everglades. Locality records
for 450 specimens are:
Broward County: Dania (1), Ft. Lauderdale (7), 16 mi. WNW, 6.2 mi. S Ft.
Lauderdale (1), 1 mi. E, 1 mi. N Hallandale (1), Hollywood (1), 1.6 mi. W Holly-
wood (3), 2.2 mi. W Hollywood (3). Collier County: 5 mi. S Deep Lake (4), Ever-
glades (64), Marco Island (3), 4 mi. W Monroe Station (1), 4.3 mi. E Ochopee
(12), Naples (3), 3 mi. SE Naples (1), 6.7 mi. SE Naples (1), 9 mi. SE Naples (1),
Royal Palm Hammock (7), 4.5 mi. SE Royal Palm Hammock (3). Dade County:
Coconut Grove (1), Coral Gables (84), Cutler (20), 2 mi. E Florida City (8), 2.2
mi. W Florida City (1), 4.1 mi. W Florida City (1), Homestead (2), 2 mi. S Kendall
(1), Lee Hammock (1), 3 mi. NW Medley (1), Miami (55), 7 mi. W, 2.5 mi. N
Miami (9), 10 mi. W Miami (1), 17.5 mi. W Miami (25), Miami Beach (24), 2 mi.
NW Miami Springs (1), Paradise Key (11), 5.4 mi. SW Paradise Key (1), 10 mi. SW
Paradise Key (1), 11 mi. SW Paradise Key (3), 20 mi. N Paradise Key (1), Rich-
mond Air Base (6), South Miami (4), 2 mi. W South Miami (1). Monroe County:
Big Pine Key (31), Cudjoe Key (2), Pinecrest (1), Snake Bight (11), Sugar Loaf Key
(25).
Eleutherodactylus ricordi planirostris Cope
Twenty males from the vicinity of Miami have snout-vent lengths
from 15.0 to 17.5 (16.6) mm.; 20 females vary from 19.5 to 25.0 (22.6)
mm. The largest specimen examined, a female from Cudjoe Key,
has a snout-vent length of 26.5 mm. Three hatchlings collected in
Miami on 25 June 1953, vary in snout-vent length from 5.5 to 6.0
(5.8) mm. The percentage of individuals with striped and mottled
color patterns has been determined (table 4). The number of speci-
mens now available more than doubles the number used by Goin for
determining color pattern variation in southern Florida specimens.
However, the ratio of striped to mottled individuals is close to that
given by him (1947: 30). The populations on the keys have the high-
est frequency of striped individuals; on the upper keys the ratio
approximates 1:1.
Essentially terrestrial in its habits, Eleutherodactylus occurs
abundantly in the mesophytic hammocks of the eastern rim of the
mainland and upper keys. On the lower keys it is found in humid


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238 BULLETIN FLORIDA STATE MUSEUM Vol. 3

sites and in edificarian situations. Calling males have been heard on
warm rainy nights throughout the summer months. No eggs of this
species were found.
TABLE 4
FREQUENCIES OF DORSAL COLOR PATTERNS IN Eleutherodactylus ricordi plani-
rostris FROM SOUTHERN FLORIDA.
Area Striped Mottled Total

Miami 23 (28.4%) 58 (71.6%) 81
Upper keys 15 (44.1%) 19 (55.9%) 34
L Lower keys 39 (40.6%) 57 (59.4%) 96


Stomach contents included largely ants (Formicidae), small beetles,
spiders, and a copepod.
Eleutherodactylus is known in southern Florida from the eastern
rim of the mainland and throughout the chain of Florida Keys. The
record given below for Cape Sable is questionable. Locality records
for 312 specimens are:
Broward County: Birch State Park (1), Ft. Lauderdale (4), Hollywood (6).
Dade County: Biscayne Bay (1), Coconut Grove (3), Coral Gables (41), Elliot Key
(6), Hialeah (2), Homestead (2), 2 mi. E Homestead (1), Lemon City (4), Matheson
Hammock (45), Miami (34), 7 mi. W, 2.5 mi. N Miami (1), Paradise Key (5), 20 mi.
N Paradise Key (1). Monroe County: Big Pine Key (18), Cape Sable (1), Cudjoe
Key (6), Islamorada, Upper Matecumbe Key (1), Key Largo (21), Key West (76),
Little Torch Key (2), Stock Island (24), Sugarloaf Key (4), Summerland Key (2).

Acris gryllus dorsalis Harlan
Females are slightly larger than males; 21 females have snout-
vent lengths varying from 18.7 to 23.9 (20.2) mm., and 10 males from
15.9 to 20.3 (18.2) mm. Apparently, this race attains a larger size
in southern Florida than it does in the northern part of the peninsula,
for Netting and Goin (1945: 304) remark that females of dorsalis
seldom exceeded 20 mm.
There is considerable variation in color pattern. Often the dor-
sal ground color is green with dark olive-green to brown bars on
the dorsolateral surfaces of the body; usually the middorsal light
stripe is well defined. However, some individuals are so darkly col-
ored as virtually to obscure the dark bars and leave no trace of the
middorsal stripe. The diagnostic thigh pattern described by Netting
and Goin is constant in specimens from southern Florida.
Acris is found in nearly all aquatic habitats in southern Florida
except salt marshes. It has been detected calling from canals and




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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 239

sloughs in the Everglades, cypress ponds, temporary pools in pine-
lands, artificial ponds, and brackish-water marshes. Singing males
are first heard in mid-March; actual choruses are formed by April
and continue until middle or late October, reaching a peak in May
and June. Recently transformed young have been collected in July
and August.
The food items contained in stomachs of 36 individuals include:
Orthoptera, Gryllidae (1); Coleoptera, Carabidae (1), Dytiscidae
(many), Hydrophilidae (many), Pselaphidae (1), Scarabaeidae (1),
Staphylinidae (1); Hemiptera, Corixidae (1); Hymenoptera, Formici-
dae (many); Diptera, Ichneumonidae (2). Aside from the above in-
sects one lycosid spider was found. Beetles comprised the greatest
percentage of bulk, followed by ants.
Acris has been removed from the stomachs of Scaphiopus, Natrix
sipedon pictiventris, and Thamnophis sauritus.
In southern Florida Acris occurs everywhere on the mainland
where there is fresh water; it reaches its greatest abundance in the
wet prairies and cypress ponds. It does not occur on the keys. The
type specimen of Acris acheta Baird, of unknown provenance, was
stated to have originated from Key West. This we believe to be in
error. Locality records for 80 specimens are:
Dade County: Coral Gables (1), Homestead (2), 4 mi. S Homestead (2), 9 mi.
N, 6.6 mi. W Homestead (1), Miami (1), Paradise Key (3), 15.9 mi. SW Paradise
Key (2). Collier County: 8.2 mi. W Monroe Station (15), Naples (11), 3 mi. SE
Naples (2), 4.2 mi. SE Naples (2), 5 mi. SE Naples (2), 7.7 mi. SE Naples (1), 4.6
mi. NW Royal Palm Hammock (12), 1.6 mi. NW Royal Palm Hammock (3).
Monroe County: 6.1 mi. S, 6.4 mi. E of Monroe Station (2), Pinecrest (18).

Pseudacris nigrita verrucosa Cope
A series of twenty males from the eastern edge of the Everglades
in Dade County displays the following variation in measurements:
snout-vent length 24.9-28.7 (26.7) mm., tibia length 12.2-13.6 (12.9)
mm., head width 7.6-8.4 (8.0) mm.; six females display the following
variation: snout-vent length 26.8-29.8 (28.2) mm., tibia length 13.5-
13.9 (13.8) mm., head width 7.8-8.8 (8.4) mm. The dorsal color pat-
tern, consisting of dark olive-green to brownish-black spots on a
green or greenish-tan ground color, is variable in the relative amount
of the dorsum covered by the spots. In some individuals the squarish
spots are large and close together so that the ground color is repre-
sented solely by narrow lines between the spots; the other extreme
has a few scattered dark spots on the dorsum, thus leaving large areas
of ground color.


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Pseudacris has been reported as breeding from mid-January to
early September in southern Florida. It appears to reach its peak
in June and July. The males call from the bases of, or in the midst
of clumps of, grasses, from the ground, or from holes and cracks in
the limestone borders of sink holes. Brady and Harper (1935: 109)
reported a clutch of 160 eggs deposited by a recently collected
female.
Examination of the stomach contents of ten individuals revealed
the remains of ants and small beetles.
Wherever Pseudacris has been found in southern Florida it has
been associated with limestone sink holes, especially bordering the
wet prairies. In the rather narrow rocky pineland-Everglades eco-
tone between Miami and Homestead the species is especially abund-
ant. Similar ecological conditions are associated with its presence
in the vicinity of Naples, Collier County. It does not occur in sandy
country, in the wet prairies proper, nor on the keys. Locality records
for 92 specimens are:
Collier County: 2.1 mi. N Deep Lake (1), 1 mi. E Naples (5), 3.1 mi. SE
Naples (1). Dade County: 4.3 mi. W Florida City (2), 9 mi. N, 2 mi. W Home-
stead (40), 9 mi. N, 7 mi. W Homestead (11), Miami (1), 19 mi. W, 6.4 mi. S
Miami (7), 19 mi. W, 8 mi. S Miami (13), Paradise Key (6), 1.3 mi. N, 5.2 mi. E
Paradise Key (5).
Limnaoedus ocularis Daudin
Hyla ocularis Daudin, 1801, in Sonnini and Latreille, Histoire Naturelle Reptiles,
vol. 2, p. 187.
Hylodes ocularis Holbrook, 1838, North American Herpetology, ed. 1, vol. 3, p. 79,
pl. 14.
Limnaoedus ocularis, Mittleman and List, 1953, Copeia, no. 2, p. 83.
Twenty males have snout-vent lengths from 13.0 to 15.5 (14.2) mm.
and tibia lengths from 7.0 to 9.0 (7.7) mm.; four females have snout-
vent lengths from 15.3 to 16.0 (15.6) mm. and tibia lengths from 7.8
to 8.1 (8.0) mm. There is little variation in coloration; the dorsal
ground color may be light yellowish tan, greenish gray, or light red-
dish brown, with or without a darker middorsal longitudinal stripe.
Limnaoedus is abundant in the wet prairies, where breeding
choruses can be detected throughout the summer. Calling males
cling to sawgrass, needle grass, or cattails and perch above the water,
sometimes at a height of three or more feet. Clasping pairs have
been found on plants above the water.
The contents of ten stomachs were examined; only the remains
of small ants, a small spider, and a small unidentified crustacean
were found.


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 241

This small species apparently ranges throughout the prairie re-
gions of southern Florida; in areas where there is an interdigitation
of prairie and pineland it is found in the pineland. A specimen of
this species is included in a small collection made for the Academy
of Natural Sciences of Philadelphia by Mr. Hebard and is purported
to be from Key West. Several forms in Hebard's collection from
Key West are unknown from the island. On the basis of the question-
able provenance of the specimens and because of the lack of appro-
priate habitat for Limnaoedus on Key West, we do not consider the
record to be valid. Locality records for 64 specimens are:
Broward County: 18.8 mi. WNW Ft. Lauderdale (13). Collier County: 2.1
mi. N Deep Lake (1), 3 mi. SE Naples (1), 4.6 mi. NW Royal Palm Hammock (1),
5.2 mi. NW Royal Palm Hammock (1). Dade County: Homestead (2), 9 mi. N,
7 mi. W Homestead (5), 7 mi. W, 2.5 mi. N Miami (18), 19 mi. W Miami (2),
6.5 mi. SW Paradise Key (13). Monroe County: 8.2 mi. W Monroe Station (7).

Hyla cinerea Schneider
Calamita cinereus Schneider, 1799, Historiae Amphibiorum, vol. 1, p. 174.
Hyla semifasciata Hallowell, 1856, Proc. Acad. Nat. Sci. Philadelphia, vol. 8, p. 307.
Hyla evittata Miller, 1899, Proc. Biol. Soc. Washington, vol. 13, p. 75.
Hyla cinerea evittata, Dunn., 1937, p. 10.
Twenty males from the vicinity of Miami display a variation in
snout-vent length from 40.2 to 50.7 (45.6) mm., three females, 50.0 to
57.5 (53.5) mm. Individuals on the keys and the eastern rim are
larger than those in the Everglades; specimens from the keys more
closely approach those from northern Florida in size (table 5).
The finding of several specimens that had incomplete or no lat-
eral light stripes prompted an investigation of the frequency of
occurrence of these color pattern variations in other parts of the
range with special attention to the validity of the subspecies evittata.
Those individuals in which the light stripe continues posterior to the
sacrum are considered as having a complete stripe; those in which
the stripe terminates between the insertion of the forelimb and the
sacrum are categorized as having a half stripe. The remaining in-
dividuals either have no stripe at all or a thin light line along the
upper lip; in these the stripe is considered to be absent. Four hun-
dred and sixty-six specimens from the southern United States were
examined (table 6). None of the samples was made up of individuals
with only one kind of color pattern, and all samples contained some
individuals with half stripes. The populations with the highest fre-
quencies of the three kinds of color pattern are: complete stripes-
Gainesville, Florida, 85 percent; half stripes-Melville, Louisiana, 70










BULLETIN FLORIDA STATE MUSEUM


TABLE 5
GEOGRAPHICAL VARIATION IN MEAN MEASUREMENTS AND PROPORTIONS OF
Hyla cinerea IN PENINSULAR FLORIDA.

Locality N Body Tibia Tibia length Head Head width
length length Body length width Body length

Silver Springs, 10 50.6 25.5 50.4% 14.7 29.1%
Marion County
Miami, 20 45.6 22.8 50.0 13.0 28.5
Dade County
Ochopee, 20 38.4 18.9 49.2 10.6 27.6
Collier County
Cudjoe Key, 12 48.0 23.3 48.5 14.4 30.0
Monroe County



TABLE 6
FREQUENCIES OF STRIPED COLOR PATTERN IN Hyla cinerea FROM
SOUTHERN UNITED STATES.

Lateral Stripe
Locality Complete Incomplete Absent Total

Florida
Big Pine Key 0 2 3 5
Cudjoe Key 4 6 3 13
Englewood 18 2 1 21
Gainesville 22 4 0 26
Miami 54 11 4 69
Ochopee 53 4 12 69
Silver Springs 17 11 3 31
Georgia
Okefenokee Swamp 5 3 1 9
South Carolina
Charleston 24 8 2 34
Virginia
Jamestown 12 3 3 18
Louisiana
Alexandria 4 1 1 6
Holden 7 2 0 9
Melville 9 36 6 51
Texas
Liberty County 7 11 3 21
Rockport 16 30 35 81
Illinois
Wolf Lake 1 1 1 3

TOTAL 253 135 78 466


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 243

percent; no stripes-Big Pine Key, Florida, 60 percent. The Big
Pine Key sample is small (5 specimens); however, of 81 individuals
from Rockport, Texas, 43 percent lacked stripes, and 37 percent had
half stripes.
Dunn (1937) in discussing color pattern variation in Hyla cinerea
in the northeastern part of its range stated: "81% in the upper tide-
water Potomac area have no stripe or a short stripe; 41% in other
parts of Maryland and Virginia have no stripe or a short one. Caro-
lina material available to me is not very extensive, but it would seem
that there only 25% have no stripe or a short stripe, whereas 75%
have a long stripe. Reports from further south indicate that 100%
long stripe occurs in the far south, especially on the Gulf Coast."
The acquisition of large series from the far south provides evidence
contradicting Dunn's statement about the frogs in that region. Noble
and Hassler (1936) and Dunn (1937) have shown that there are no
morphological differences correlated with the different color-pattern
types.
From the material now available it is clearly demonstrated that
the "normal" striped and the evittate, as well as all degrees of inter-
mediacy, are found throughout the range of the species. In some
areas the frequency of evittate individuals is high, but when these
occur in such distant areas as Maryland and coastal Texas, the pres-
ence or absence of a lateral stripe loses significance as a systematic
character. Consequently, Hyla evittata Miller should be considered
a strict synonym of Hyla cinerea Schneider.
Hyla cinerea breeds in southern Florida from mid-May to early
October. Calling males are usually found on bushes and trees,
seldom on the ground. They have been observed about 15 feet above
the ground in palm trees. Daytime retreats include axils of palm
fronds and shaded branches in hammocks.
Food items found in stomachs of ten individuals included 2
elaterid and 2 scarabaeid beetles, 1 lepidopteran larva, 1 chalcid,
and 12 termites.
Hyla cinerea is found throughout southern Florida and the keys
with the exception of mangrove swamps and salt marshes. It is par-
ticularly abundant in willow clumps along canals and sloughs, in
mesophytic hammocks, and in cypress swamp. It is least abundant
in the sandy pineland. Locality records for 455 specimens are:
Broward County: 2 mi. W Davie (2), 8 mi. W, 5 mi. S Hallandale (1), Holly-
wood (9), 10 mi. S Pompano (1). Collier County: 2.5 mi. E Carnestown (5), 1.3 mi.
WNW Collier-Dade Co. line (2), Deep Lake (2), 2.1 mi. N Deep Lake (8), Ever-
glades (1), Marco Island (7), 2.9 mi. W Monroe Station (28), 8.6 mi. E Monroe


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Station (6), Naples (3), 3-5 mi. SE Naples (19), 7-9.4 mi SE Naples (19), 4.2 mi. N
Naples (3), 4.3 mi. E Ochopee (53), 2.5 mi. W Ochopee (3), 5 mi. W Ochopee (1),
Royal Palm Hammock (11), Tamiami Trail (1), Turner River (3). Dade County:
Coconut Grove (1), Coral Gables (2), 14 mi. W Coral Gables (1), 16 mi. SW Coral
Gables (2), 4.2 mi. S Florida City (3), 5.2 mi. E Florida City (1), Homestead (13),
Lemon City (12), Little River (1), Matheson Hammock (1), Miami (47), 7 mi. W,
2.5 mi. N Miami (7), 8.2 mi. W Miami (13), 16.9 mi. W Miami (57), 18 mi. W
Miami (5), 22.7 mi. W Miami (1), Medley (1), Paradise Key (33), 6.5 mi. SW
Paradise Key (12), 9.3 mi. N, 5.2 mi. E Paradise Key (1), 20 mi. N Paradise Key
(2), Perrine (1), South Miami (1). Monroe County: Big Pine Key (9), Cudjoe Key
(13), Flamingo (7), Islamorada, Upper Matecumbe Key (1), Matecumbe Key (1),
5.5 mi. S, 6.3 mi. E Monroe Station (2), 50 mi. W Miami (1), Pinecrest (11).

Hyla femoralis Bose
The largest specimen examined is a female with a snout-vent
length of 39 mm. Thirty-eight males have an average snout-vent
length of 80.7 mm., tibia length of 14.8 mm., and snout-vent/tibia
ratio of 45.8 percent; 19 females average 34.2 mm., 15.6 mm., and
45.6 percent. Usually there is a dark brown stripe separating the
reddish dorsal ground color from the white venter. In specimens
from southern Florida this stripe is distinct in most, but in 25 percent
it is poorly defined, and in 3 percent, it is absent. Individuals from
northern Florida (Silver Springs) are much like southern specimens
in this respect. In a series of 22 males from Charleston, South Caro-
lina, only 27 percent have a distinct stripe, whereas the stripe is
poorly defined in 50 percent and absent in 23 percent. Some indi-
viduals were greenish gray or pale green when found. One from
Marco Island is pale greenish gray with little suggestion of a dorsal
pattern, consequently giving the appearance of Hyla squirella. Al-
though we have not observed individuals changing color, Wright and
Wright (1949: 323) intimate that the species exhibits metachrosis.
For the most part Hyla femoralis is associated with pine forests;
two specimens have been found in the rosemary scrub association on
Marco Island. It lives in the sandy areas and not in the rocky pine-
land.
In southern Florida breeding takes place from June to early
October. A large chorus was encountered 3 miles southeast of
Naples on 5 July 1953. The males were calling from vines, grasses,
and shrubs to a height of about two feet above the water in a tem-
porary pond in the pine flatwoods. Clasping pairs were found on
boles in the pines, and other pairs were seen higher on the trees than
one could reach. Drainage in the pine flatwoods is rapid, and a heavy
rain must fall before any appreciable amount of standing water can


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 245

be found. Consequently, breeding choruses of Hyla femoralis are
likely to be heard only after heavy rains.
The stomachs of 20 individuals were examined for food remains;
of these one was empty, and two contained only plant material (bark
and plant buds). The others contained mostly insect remains, as fol-
lows: 4 grasshoppers (Tetrigidae), 1 cricket (Gryllidae), 6 beetles (Car-
abidae, Elateridae, Scarabaeidae), 1 caddisfly, 2 ants (Formicidae),
4 wasps (Vespidae), 3 unidentified insects, and 1 jumping spider
(Attidae).
Hyla femoralis just enters the area of study; in peninsular Florida
its range terminates in the sandy pine flatwoods of southeastern
Broward County and western Collier County. It does not occur in
the cypress forests nor in the Everglades. Locality records for 63
specimens are:
Broward County: 1 mi. N Hallandale (1), 10 mi. S Pompano (3). Collier
County: Marco Island (2), 3 mi. SE Naples (41), 4.6 mi. NW Royal Palm Ham-
mock (16).
Hyla gratiosa Le Conte
Twenty-six males from Collier County vary in snout-vent length
from 54.4 to 70.3 (61.5) mm.; two females have snout-vent lengths
of 56.8 and 58.2. Examination and comparison of 30 specimens
from southern Florida with 45 individuals from Gainesville and 27
from South Carolina shows that specimens from southern Florida
resemble those from South Carolina most closely in size; specimens
from Gainesville are slightly smaller. Specimens from the western
part of southern Florida, however, are at once distinguishable from
individuals from South Carolina on differences in color pattern.
Specimens from North Carolina, South Carolina, Georgia, and Lou-
isiana are characterized by having a prominent white line along the
upper lip posterior to the tympanum, a distinct white line along the
outer margin of the forearm, a definite white line dorsal to the vent,
and a yellowish line beginning at the shoulder and proceeding pos-
teriorly along the side. Specimens from southern Florida (exclusive
of Broward County) have the lip line absent or almost so due to en-
croachment of pigments from above the lip, the white line along
the forearm absent or almost so, white pigmentation above the vent
and in the groin scattered and not forming a white bar, and the yel-
lowish lateral lines indistinct and blending quickly into the lateral
pattern of spots, or represented only by a short lateral bar immedi-
ately behind the shoulder. Specimens from Broward County seem
typical of Hyla gratiosa as it occurs in South Carolina and Georgia.


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This species has been taken only in the vicinity of ponds, usually
with cypress as the predominant tree, but occasionally in low ponds
in flatwoods where cypress was absent. These ponds are located in
sandy pineland, where Pinus clausa occurs. Calling males have been
heard from mid-June through September. Usually males call while
floating in water with the head and vocal sac above the surface and
the legs distended posteriorly. Such aquatic males take alarm easily
and often escape capture. One calling male was taken from the top
of a three-foot high cypress stump, and two silent males were taken
from boles of living cypress trees. Clasping pairs were collected in
the water. When calling from the water, the males usually were in
open areas away from vegetation; however, occasionally a male was
heard calling from dense cover.
The stomachs of the individuals examined were empty.
Hyla gratiosa previously has not been reported from southern
Florida. Carr (1940: 60) listed Hillsborough County on the west coast
and Saint Lucie County on the east coast as the southernmost locali-
ties where this frog had been taken. Also he listed no counties in the
central part of the state to the south of Marion and Osceola counties.
Apparently H. gratiosa extends southward along the coastal areas,
and the southern populations are separated by the broad expanse of
the Everglades and the more northern prairies. The differences
pointed out between individuals from Broward County and those
from the western counties are probably related to the absence of
genetic continuity between these terminal populations, except via
breeding populations to the north along the peninsula. Locality
records for 32 specimens are:
Broward County: No additional data (2), 10 mi. S Pompano (2). Collier
County: 2.1 mi. N Deep Lake (2), 3 mi. SE Naples (16), 4.2 mi. N Naples (8),
13.6 mi. NW Royal Palm Hammock (2).

Hyla septentrionalis Boulenger
Barbour (1931b) first reported the presence of this species in
Florida; he mentioned that old residents of Key West recalled that
the frogs had always been there. Hyla septentrionalis is widely dis-
tributed in the Bahama Islands and Cuba; it has relatives throughout
the Antilles. Obviously this group of frogs is capable of insular dis-
persal, so there is little reason to consider Hyla septentrionalis as a
species introduced into the fauna of the United States. Schwartz
(1952b) first reported the species from the mainland.
In size the females are much larger than males; the largest fe-
male examined has a snout-vent length of 96.5 mm.; the largest male,


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61.6 mm. There is little difference in the size and proportions of
individuals from Key West and those from Miami (table 7). In life
the dorsal ground color changes from light green to ashy gray to light
brown, sometimes to reddish brown. The dorsal markings are dark
gray to dark olive green or deep brown. Usually the markings form
reticulations, but this is highly variable (fig. 21).

TABLE 7
MEAN MEASUREMENTS AND PROPORTIONS OF Hyla septentrionalis
IN soTrHERN FLORIDA.

Locality Sex N Body Tibia Tibia length Head Head width
length length Body length width Body length

Miami a 20 57.5 26.5 46.2% 17.7 30.7%
9 7 71.7 34.1 47.6 23.3 33.2
Key West S 20 53.0 24.0 45.3 16.6 32.9
9 10 78.6 36.1 46.1% 26.1 33.3%


Figure 21.-Variation in the dorsal color pattern in Hyla septentrionalis
from Miami.

Calling males have been encountered from early March to early
September. On 7 September egg masses were observed in an outdoor
aquarium in Miami; these formed a thin sheet on the surface of the
water. A captive female laid 130 eggs in an aquarium. Twenty-four
hours after deposition the average diameter of the outer envelope (10
eggs) was 3.0 mm.; that of the inside envelope 2.4 mm.; that of the


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embryo 2.0 mm. Shortly before hatching the largest embryo was 2.8
mm. in length. Eggs laid on 10 July hatched 27V2 hours after deposi-
tion. Forty-eight hours after the eggs were laid the tadpoles aver-
aged 2.1 mm. in snout-vent length and 3.8 mm. in tail length. The
abdominal region was swollen with yolk; the eyes were covered,
and stout, rounded gills were present (fig. 22). At an age of five days
the tadpoles had a snout-vent length of 3.9 mm. and a tail length of
5.1 mm.; at seven days these increased to 4.2 and 6.0 mm., and at
twenty days to 6.0 and 8.4 mm. At 54 days the average snout-vent
length was 12.6 mm. and tail length 19.6 mm. No limbs had appeared
at this time. The tadpoles have a round body and a wide caudal fin.
They are predominantly black above; the fleshy part of the tail is
grayish brown; the fin is transparent with scattered melanophores
(fig. 23). The teeth are arranged in six rows: one complete and one
incomplete row above the horny beak, and three complete and one
incomplete rows below the beak (fig. 24). Sixteen recently trans-
formed young collected on 25 July 1952, have an average snout-vent
length of 12.6 mm.; the tibia/snout-vent ratio is 0.50, and the head
width/snout-vent ratio is 0.33. The young have a ground color that
is pale green to light tan with a rather broad dorsolateral cream to
yellow stripe. The young individuals were found clinging to leaves
of bushes and blades of grass in and around a pond.
Peterson, Garrett, and Lantz (1952) reported collecting breeding
individuals of Hyla septentrionalis from brackish water on Key Vaca.
If the eggs and tadpoles of this species are capable of development
in brackish water, since fresh water is lacking on some islands, this
ability would be helpful in the dispersal of the species throughout
the chain of Florida Keys.
Stomach contents of twenty individuals from Miami revealed an
assortment of arthropod remains: 5 beetles (Curculionidae, Scarabaei-
dae, Elateridae), 1 roach (Blattidae), 1 cricket (Acrididae), 6 bugs
(Cercopidae, Scutelleridae), 2 lepidopteran larvae, 1 may-fly, 24 pill
bugs, and a small crustacean.
Hyla septentrionalis is an inhabitant of mesophytic and edificarian
situations. It is unknown from the prairie and pineland habitats.
At Key West the frogs are commonly found in old cisterns, and a resi-
dent informed us that they are sometimes found in toilet bowls!
These frogs are distributed over the Florida Keys and are abundant
in certain areas in Miami. Allen and Neill (1953) mentioned the
presence of this species at Paradise Key, where it had been released
some years before. No specimens are available from that locality.
Locality records for 227 specimens are:


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Figure 22.-Forty-eight hour tadpole of Hyla septentrionals. Total length
5.9 mm.










Figure 23.-Fifty-four day tadpole of Hyla septentrionalis. Total length
32.2 mm.


Figure 24.-Mouthparts of fifty-four day tadpole of Hyla septentrionalis.


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BULLETIN FLORIDA STATE MUSEUM


Dade County: Miami (45). Monroe County: Big Pine Key (1), Key Largo
(1), Key Vaca (35), Key West (102), Matecumbe Key (2), Stock Island (22), Upper
Matecumbe Key (19).

Hyla squirella Bosc

The dorsal ground color of specimens from southern Florida may
be either unicolor greenish gray or light brown, or variously mottled
or spotted with dark greenish or brown (depending upon the ground
color of the individual); often there is an interocular dark bar. Sixty
adult males have snout-vent lengths from 24.2 to 43.0 (32.6) mm., 21
females, 23.3 to 35.1 (29.0) mm. Comparison of four series of H.
squirella from various localities indicates that specimens from south-
ern Florida show striking variation (table 8). Whenever possible,
the 15 largest individuals of each sex from each area were measured.
A series from Collier County, when compared with specimens from
Miami, and specimens from Charleston County, South Carolina, and
Marion County, Florida, shows that the populations west of the Ever-
glades are noticeably smaller in the measurements taken. The series
from Miami is comparable in all measurements (although there are
certain slight average differences) to specimens from northern Florida
(Marion County) and South Carolina. Average differences in the
size of the sexes is not striking.

TABLE 8
AVERAGE MEASUREMENTS OF FOUR SAMPLES OF Hyla squirella FROM
SOUTHEASTERN UNITED STATES.

Locality Sex N Body Tibia Foot Head Head Interorbital
length length length width length distance

Charleston, 8 15 33.0 12.9 12.6 10.5 9.4 3.1
South Carolina 9 6 33.4 12.7 11.8 10.2 9.0 2.8
Marion County, 8 15 31.6 12.6 13.0 10.9 9.3 3.1
Florida 9 15 33.4 13.2 13.6 11.1 10.0 2.9
Miami, Florida 8 15 33.9 12.0 11.6 10.9 9.3 2.8
9 7 31.5 11.2 11.2 9.7 8.7 2.8
Collier County, $ 15 26.6 9.3 9.1 8.1 7.3 2.2
Florida 9 14 27.8 ,10.4 10.8 9.0 8.6 2.3


Material from the Florida Keys is not abundant. Fifteen adult
males from the lower keys have an average snout-vent length of
32.9 mm., and thus are more like the specimens from Miami. A


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 251

series of 15 males from Miami Beach have an average snout-vent
length of 36.9 mm., with the largest being 43.0 mm.; these are the
largest specimens of H. squirella examined. The discrepency in size
between Miami, Miami Beach, and Collier County individuals is
noticeable in the field; these differences are even more striking in the
laboratory, when males from these three localities are placed side
by side.
This frog has been taken in all major habitats in southern Florida
except mangroves; however, specimens from the vicinity of Flamingo
were collected while in chorus on a prairie adjacent to a marine
canal; and undoubtedly the frogs occupy mangroves in such areas.
When calling, this frog utilizes almost any small body of water,
but temporary rain pools of moderate depth seem to be preferred.
Vocalizing individuals often perch on debris and twigs above the
water, but they have been seen singing from the water, on the ground
near water, and hidden in clumps of grass. When calling, the frogs
create an almost unbearable din with their raucous voices, which
resemble a harsh, short, ascending scale; the noise so created is
especially distracting when it results from a chorus in a confined area,
such as a deep sink hole. Loud choruses often completely obscure
the calls of other species in the same pond. When not calling, H.
squirella has been collected in hammocks, under logs and debris, in
the palm thatching of a Seminole cheki, and upon the rails of bridges
along the Tamiami Trail on rainy nights. Numerous individuals
were found during the day beneath the bark of dead pine stumps
west of Homestead. This is the only hylid frog so encountered.
The "rain call" of H. squirella is familiar to most residents of
southern Florida. This call is uttered by lone males from trees and
bushes (usually during the day) when the weather becomes cloudy
and sultry and rainfall is impending. The call is a raucous quack
and differs from the breeding call. This species seems more sensi-
tive to pre-precipitation weather changes than most other frogs in
the area. In Miami, on one occasion, no H. squirella were heard
calling until a warm, sultry northern wind began. As the velocity
of the wind increased (without rain), the frogs began vocalizing in
gradually increasing numbers, so that by the time the rain began they
were in loud chorus. Dates for chorusing of H. squirella range from
30 March to 8 August.
Stomachs of 20 individuals were examined and the following food
items were noted. Nine stomachs were empty; two contained only
plant debris, and beetles (in two cases identified as Elateridae) were


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252 BULLETIN FLORIDA STATE MUSEUM Vol. 3

found in four stomachs. Crustacean remains (? very small crayfish)
were identified from two frogs, and remains of a spider, a cricket,
and an ant each were encountered in a single stomach.
Hyla squirella is widespread in southern Florida. It occurs
throughout the keys. Locality records for 685 specimens are:
Broward County: 1 mi. W Davie (52), 2 mi. W Davie (31), Hallandale (28),
Hollywood (48). Collier County: 1.3 mi. WNW Collier-Dade Co. line, Tamiami
Trail (5), 2.1 mi. N Deep Lake (12), Everglades (7), 8.6 mi. E Monroe Station
(41), Naples (1), 1-3 mi. SE Naples (10), 6-10 mi. SE Naples (3), Royal Palm
Hammock (40), 4.6 mi. NW Royal Palm Hammock (2), 8-13 mi. NW Royal Palm
Hammock (10), Tamiami Trail (1). Dade County: Coconut Grove (6), Coral
Gables (8), 16 mi. SW Coral Gables (1), 1.5 mi. SE Florida City (28), 4.3 mi. W
Florida City (4), Homestead (37), Lemon City (14), Long Pine Key (9), Matheson
Hammock (1), Miami (105), 7 mi. W Miami (20), 20 mi. W Miami (15), Miami
Beach (33), Opa-locka (1), Paradise Key (23), 6.5 mi. SW Paradise Key (3), 20
mi. N Paradise Key (1), Uleta (12). Monroe County: Big Pine Key (14), Boca
Chica Key (1), Coot Bay (1), Cudjoe Key (1), Flamingo (22), 1 mi. NE Flamingo
(4), Key Vaca (3), Key West (2), Lignumvitae Key (2), Little Torch Key (12), Long
Key (4), Lower Matecumbe Key (1), Matecumbe Key (5), Pinecrest (1).

Gastrophryne carolinensis carolinensis Holbrook
Engystoma carolinensis Holbrook, 1836, North American Herpetology, ed. 1, vol.
1, p. 83.
Eighty-seven males have snout-vent lengths from 18.8 to 30.5
(26.2) mm.; 49 females, from 22.4 to 32.5 (28.3) mm. The dorsal color
pattern varies from an almost unicolor tan with only a faint indication
of the dorsal blackish stripes to a dorsum with two light bands, each
edged heavily in black. The venter may, be heavily spotted or may
be tan and relatively unicolor. These differences in coloration have
led Hecht and Matalas (1946) to segregate specimens from the Florida
Keys into three categories: 1) carolinensis-dorsum dark and blotched
or with indistinct dorsolateral stripes, and venter mottled; 2) "Key
West"-dorsal pattern of two prominent light tan dorsolateral stripes
bordered by a distinct dark margin on a tan background; 3) olivacea-
like-virtually without pattern and thus with a uniformly light tan
dorsum, with ventral coloring usually much reduced. They found
(1946: 3) that 39 specimens from the keys might be divided into 9
carolinensis (23%), 19 "Key West" (48%), and 11 olivacea-like (29%).
Examination of a series of 78 other individuals from the keys shows
that 12 percent are carolinensis, 34 percent are "Key West," and 24
percent are olivacea-like. Intermediates between "Key West" and
carolinensis formed 9 percent of the sample, and intermediates be-
tween "Key West" and olivacea-like made up 21 percent. Thus of


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 253

78 specimens, only 12 percent are strictly comparable to G. c. carolin-
ensis, whereas 88 percent show other types of coloration. Populations
of Gastrophryne on the keys show differences in pigmentation and
patterning; isolation by salt water probably has been responsible in
establishing these patterns on the islands, apparently to the detriment
of the typical carolinensis pattern. A series of 48 individuals from
Miami shows that 56 percent are carolinensis, 16 percent are "Key
West," 2 percent are olivacea-like, and 25 percent are intermediate
between "Key West" and carolinensis. Thus, on the southern Florida
mainland the population still shows the mixed character which is
noted on the keys, although the preponderance of specimens shows
a carolinensis pattern. Finally, 86 specimens from South Carolina
show 86 percent carolinensis characters, and 14 percent (5 specimens)
are variously intermediate.
Gastrophryne is common around human dwellings, often being
taken under trash and boards in the vicinity of houses during the day.
A single individual was taken from the house of a wood rat (Neotoma
floridana) on Key Largo. Other than under debris, this frog is seldom
seen during the day; small individuals were seen abroad on two oc-
casions.
The call is a sheeplike "baaa" which, when accompanied by large
choruses of other amphibians, may be virtually inaudible in the gen-
eral din. Choruses exclusively of Gastrophryne are seldom heard.
The shy males call from sheltered situations, 'often from beneath
trash at the edge of the water or partially buried in grass. The call
is given with the head protruding above the surface and with the hind
legs widespread as the frog lies extended in the water. At times the
males give voice on the bank away from the pond; in instances when
these individuals have been found, they were located in small de-
pressions filled with water. Calling males have been collected be-
tween 27 April and 9 July, and presumably breeding takes place
within this period.
Examination of several stomachs shows that the diet of these frogs
consists solely of ants. The taking of an individual in an ant nest
in Tennessee has been mentioned by Wood (1948).
Gastrophryne occurs throughout southern Florida on the mainland
and on the keys. Locality records for 278 specimens are:

Broward County: 2 mi. W Davie (5), Ft. Lauderdale (1), 6 mi. W Ft. Lauder-
dale (3), 2 mi. S junct. Ft. Lauderdale road and Okeechobee-Miami road (1),
Hallandale (2), Hollywood (10). Collier County: Everglades (2), 3.7 mi. SE Naples
(1), 7.5 mi. SE Naples (2), 2.8 mi. S Miles City (1). Dade County: Coral Gables


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254 BULLETIN FLORIDA STATE MUSEUM Vol. 3

(1), 5 mi. E Florida City (1), 1.5 mi. SE Florida City (26), Hialeah (1), Home-
stead (2), 6 mi. S Homestead (2), Miami (56), 7 mi. W, 2.5 mi. N Miami (8), 17
mi. W Miami (6), 19 mi. W, 3 mi. S Miami (1), Miami Beach (3), Miami Springs
(2), Paradise Key (3), 5.2 mi. NE Paradise Key (1), 9 mi. W Paradise Key (3), Uleta
(3). Monroe County: Big Pine Key (9), Coot Bay (2), Cudjoe Key (1), Key Largo
(1), Key West (82), Lignumvitate Key (8), Little Torch Key (2), Matecumbe Key
(7), Pinecrest (3), Stock Island (9), Sugarloaf Key (2), Summerland Key (5).

Rana areolata aesopus Cope
Rana areolata aesopus Cope, 1886, Proc. Amer. Phil. Soc., vol. 23, p. 517. Neill,
1957, Herpetologica, vol. 13, pt. 1, p. 52.
On 19 April 1957, Dennis R. Paulson collected one individual of
this species in a cypress bottom in the relatively dry scrub area near
North Naples, Collier County. Others have been heard calling in
the vicinity of Naples. That this locality is not extremely disjunct
from the rest of the range is shown by records of the species in Char-
lotte County (L. Neil Bell, in litt.) and Okeechobee County (D. R.
Paulson, in litt.). The specimen from North Naples (DRP 500) is a
male with a snout-vent length of 95 mm. and constitutes the only
locality record.
Rana grylio Stejneger
Thirteen adult males have snout-vent lengths from 96.7 to 117.0
(106.0) mm., 12 females, from 98.1 to 135.6 (115.2) mm. The tibia/
snout-vent length ratio is greater in males (46.1% as opposed to 44.1%
for females); the diameter of the tympanum/head width ratio is 41.6
percent in males and 27.2 percent in females. The dorsal ground
color varies from pale green to grayish green or light olive green.
The dorsal markings are dark olive green to brown or black. The
belly is creamy white; the undersurfaces of the hind limbs are boldly
mottled with dark gray or black. Usually there are some dark spots
on the abdomen and a large dark area in the thoracic region. In
some there is a gray and cream mottling over the entire ventral sur-
faces (fig. 25). In comparison with specimens from northern Florida
and southern Mississippi, individuals from southern Florida seem to
have less brown pigment in the dorsal coloration and to have a more
boldly marked venter.
Six juveniles collected on 18 May 1952, have an average snout-
vent length of 36.6 mm. Dorsally they are colored like adults, but
the ventral surfaces are cream with dark chocolate-brown mottling.
Rana grylio occurs most commonly in the wet prairies and occa-
sionally in sloughs. The optimum habitat appears to be moderately


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 255
open prairie with a subsurface mat of vegetation upon which to rest.
Juveniles were collected in masses of water hyacinths.



10 _


Figure 25.-Variation in the ventral markings of Rana grylio from the Ever-
glades in Dade County.

The piglike grunts of the males are heard throughout the year;
however, even at the height of the breeding season, apparently in
June and July, choruses are not encountered. Individual males call
from scattered spots in the prairie. Tadpoles have been observed
in sloughs that were choked with water hyacinths.
Examination of food remains in the stomachs of ten adults re-
vealed: 2 beetles (Dystiscidae and Scarabaeidae), 5 bugs (Belostomati-
dae), 6 wasps (Vespidae), 2 grasshoppers (Acrididae), 2 spiders (Ly-
cosidae), and 1 crayfish (Procambarus alleni). Stomachs from eight





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I


juvenile frogs contained: 14 aquatic beetles (Dystiscidae, Haliplidae,
and Hydrophilidae), 1 bug (Naucoridae), 1 earwig (Forficulidae),
1 spittle-bug (Cercopidae), 1 chalcid (Chalcididae), and 2 spiders
(Lycosidae). Data on the feeding habits of Rana grylio supplied by
the Game and Fresh Water Fish Commission of the State of Florida
indicate that the principal foods are: crayfish (Procambarus alleni)
very frequent, leeches (Placobdella rugosa) occasional, aquatic Hemip-
tera and Coleoptera frequent, spiders frequent, fishes (primarily small
cyprinodonts and poeciliids) frequent, and occasional small verte-
brates (Hyla cinerea, Rana pipiens, juvenile Rana grylio, and Natrix
sipedon). Carr (1940: 67) stated that crayfish were present in about
40 percent of the many specimens he examined.
The value of Rana grylio as a food item for man has been the
cause for reduced size of populations in southern Florida. Frogging
is highly commercialized, and large numbers of this species and
Rana pipiens are secured by gigging from air boats at night.
Rana grylio is widely distributed in the Everglades but does not
occur in the pinelands nor on the keys. Locality records for 62
specimens are:
Broward County: Hollywood (3). Collier County: 2-4 mi. NW Carnestown
(2), 2.7 mi. S Miles City (1), 6 mi. E Monroe Station (1), 10.3 mi. E Monroe Sta-
tion (2), 5 mi. SE Naples (1), 9.4 mi. SE Naples (2), 11.1 mi. SE Naples (2), 7.2
mi. E Ochopee (1), 7.1 mi. W Ochopee (1), 12.1 mi. W Ochopee (1), 18.9 mi. W
Ochopee (1), 10-12 mi. E Royal Palm Hammock (2), 11 mi. W Royal Palm Ham-
mock (2). Dade County: Homestead (1), Lemon City (1), Miami (5), 6 mi. W
Miami (1), 17-25 mi. W Miami (16), 40 mi. W, 2.6 mi. NW Miami (1), Paradise
Key (7), 10 mi. SW Paradise Key (1). Monroe County: 45 mi. W Miami (1),
Pinecrest (6).
Rana pipiens Schreber
Rana pipiens Schreber, 1782, Naturforscher, vol. 18, p. 185, pl. 4.
Rana halecina sphenocephala Cope, 1886, Proc. Amer. Phil. Soc., vol. 23, p. 517.
In specimens from the Everglades and surrounding pineland, the
dorsal color pattern typically consists of elongated dark brown spots
on a light tan or green ground color; the dorsolateral folds are prom-
inent and usually of a bright yellow color. The transverse bands on
the femur are incomplete; the posterior surfaces of the thighs and
the lip region are mottled with brown and cream or white. The un-
dersurfaces are white or cream, and the brown tympanum usually
has a central yellow spot. The above description is adequate for
most specimens from the mainland; however, those from Marco
Island, Collier County, may exhibit a slightly different coloration,
whereas those from the lower keys are strikingly different (fig. 26).


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 257


Figure 26.-Variation in the dorsal coloration of Rana pipiens. The individ-
ual on the right is from Big Pine Key, Monroe County, and is typical of the color-
ation exhibited by the population on the keys. The other two individuals are
from Hollywood, Broward County, and show the normal variation in coloration
exhibited by the population on the mainland.

Specimens from Marco Island usually are darker in over-all colora-
tion than individuals from the mainland. The dorsal ground color
is dark green; the spots often are nearly black. The belly is a dirty
white and mottled with dark brown or black in the throat region.
Series of specimens from Big Pine Key, Little Torch Key, and Key
West are very dark. The dorsal ground color is a dark brownish
black; the dorsal spots are black, and the ventral color is a grayish
to brownish cream, the throat and flanks often being heavily mottled
with dark gray or black. In the large adults the dorsolateral fold
usually is a bronze color. The light tympanic spot is present. Al-
though young individuals from the keys are darker than specimens
of a similar size from the mainland, they are not so dark as the adults.
Apparently, the dark pigmentation increases with age.
The isolated insular populations of Rana pipiens show a colora-
tion very similar to that displayed by individuals of the species from
southern Arizona. Duellman (1955b: 5) in discussing the Arizona
frogs postulated that the coloration might be an expression of environ-
mental conditions and not be genetically controlled.
The largest female from the mainland has a snout-vent length of
99.6 mm.; the largest male, 85.2 mm. The largest individuals of each
sex from the keys are from Big Pine Key and measure 97.7 mm.
(female) and 80.0 mm. (male). The smallest completely transformed


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specimen also is from Big Pine Key; it has a snout-vent length of 24
mm. There are no significant differences in measurements and pro-
portions between the series from the lower keys and the mainland.
Only the coloration seems to be consistently different.
Rana pipiens inhabits all areas of the mainland of southern Florida
where there is permanent water. Large populations thrive in the
Everglades, but the frogs also are abundant in sloughs, canals, and
rockpits. On Big Pine Key they inhabit a few large pot holes, an
abandoned well, and a roadside ditch. On Marco Island several
were found in an abandoned well, and on 3 July 1954, large numbers
were found in temporary ponds between the beach and a mangrove
swamp. In September 1954, at West Lake near Coot Bay Ranger
Station numerous small Rana pipiens were seen leaping from the
roadbed into the brackish water of the mangrove swamp. Deckert
(1922) mentioned seeing many Rana pipiens along the bank of a
brackish canal between Paradise Key and Cape Sable.
The possible adaptations of Rana pipiens to brackish water is an
interesting problem in the life history and ecology of this frog in
southern Florida. The specimens found in the vicinity of Coot Bay
are several miles from any permanent body of fresh water, for the
broad expanse of the Everglades gradually merges with the brackish
coastal prairies and mangrove swamps. At times of heavy rain and
corresponding high water in the Everglades, fresh water extends
farther toward the coastal areas than it does at time of drought. This
brings about a fluctuation of salinity and a stratification of salt and
fresh water. Although frogs have been seen submerged and swim-
ming in water that was definitely brackish, no eggs or tadpoles have
been found in brackish water. Certainly the frogs do not move in-
land to breed; even though Rana pipiens may be capable of moving
considerable distances within a relatively short period of time, it is
doubtful if the whole population moves several miles to breed! Also,
the presence of many small and less mobile frogs, such as Acris gryl-
lus, Hyla squirella, and Gastrophryne carolinensis, negates that pos-
sibility. Possibly there is a certain amount of tolerance to saline con-
ditions by the tadpoles and perhaps the eggs. A conclusion can be
reached only after laboratory experimentation.
Carr (1940: 68) stated that Rana pipiens in Florida breeds every
month of the year. Males may be heard throughout the year, but it
is doubtful if breeding activity is continuous. They have been noted
breeding in southern Florida from May to December, Deckert (1921:
21) recorded R. pipiens breeding at Lemon City on 5 December 1920,


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 259

where he found three batches of eggs partly floating and attached
to weed stems in about 12 to 15 inches of water in a rock pit. On a
rather cold day in February nearly 50 R. pipiens were found huddled
in a mass under a large board about 30 feet from a canal near Pine-
crest. They were quite lethargic and reluctant to move.
The following invertebrates were found in 24 stomachs: 2 centi-
pcdes (Scolopendridae), 4 grasshoppers (Tetrigidae), 1 roach (Blatti-
dac), 1 squash bug (Coreidae), 6 beetles (Carabidae, Scarabaeidae,
and Tenebrionidae), 4 lepidopteran larvae, 6 ants (Formicidae), 2
spiders (Lycosidae), and 5 snails (Helicinidae). One specimen from
Big Pine Key contained a Hyla squirella, and two adults from Marco
Island had eaten full grown Bufo quercicus.
Rana pipiens is found in all situations where permanent water
is present on the mainland and on the lower keys. Locality records
for 222 specimens are:
Broward County: Ft. Lauderdale (1), Hollywood (15). Collier County:
Carnestown (1), 5.1 mi. N Carnestown (1), Deep Lake (2), Everglades (1), Marco
Island (12), 3-5 mi. SE Naples (7), 8-10 mi. SE Naples (5), 2.5 mi. W Ochopee
(1), Royal Palm Hammock (1), 3.9 mi. NW Royal Palm Hammock (2), 13.6 mi. NW
Royal Palm Hammock (1), 7.9 mi. E Royal Palm Hammock (1). Dade County:
Coral Gables (1), 7 mi. W Coral Cables (1), 2 mi. E Florida City (2), Homestead
(2), 4-6 mi. S Homestead (6), 9 mi. N, 2 mi. W Iomestead (3), Lemon City (3),
Miami (31), 17-19 mi. W Miami (5), 19 mi. W, 3-4 mi. S Miami (7), 24-25 mi.
W Miami (3), 36 mi. W Miami (5), Paradise Key (5), 2.1 mi. E Paradise Key (1),
1.3 mi. N, 5.2 mi. NE Paradise Key (3), 20 mi. N Paradise Key (2), 1.5 mi. SW
Paradise Key (1), 15.9 mi. SW Paradise Key (1). Monroe County: Big Pine Key
(59), Key West (12), Little Torch Key (6), Pinecrest (11), Ramrod Key (1).

( ,. i. Itr osceola Stejncger
Chelydra osceola Stejneger, 1918, Proc. Biol. Soc. Washington, vol. 31, p. 89.
Chelydra serpentina osceola, Rust, 1934, Bliitter Aquarien-Terrarien-kunde, vol. 45,
p. 59.
Chelydra osceola, Richmond, 1958, p. 41.

No large specimens of this species were secured during our field
work in southern Florida. The largest specimen, a mature female,
has a carapace length of 240 mm., carapace width of 188 mm. and a
shell depth of 111 mm. A female with a carapace length of 155 mnm.
is not mature. The smallest juvenile has a carapace length of 28
mm., a carapace width of 29 mm., and a shell depth of 17 mm. The
characters used by Richmond (1958) to distinguish osceola from
serpentina apply equally well to our specimens from southern Florida.


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The fleshy ventral surface, especially in the juveniles, is dark ashy
gray to black with small cream flecks; in some there is fine cream mot-
tling on the undersurfaces of the limbs. The dark color of the fleshy
parts and of the plastron fades to a light gray or cream in the adults.
Chelydra has been encountered in canals, sloughs, and ponds.
Juveniles were collected by seiving water hyacinths in side canals
along the Tamiami Trail. Apparently the species is found in all fresh-
water environments in southern Florida, but it probably reaches its
greatest abundance in the canals in the Everglades. Locality records
for 23 specimens are:
Collier County: Everglades (1), Naples (1), 13.6 mi. SW Royal Palm Ham-
mock (1). Dade County: Biscayne Bay (1), Florida City (1), 4.2 mi. S Florida
City (2), 18 mi. N Homestead (1), Miami (2), 15 mi. W Miami (2), 17 mi. W Miami
(2), 19 mi. W, 3-6 mi. S Miami (3), 20-23 mi. W Miami (6).

Sternotherus odoratus Latreille
In specimens from southern Florida both the upper and lower
beaks are black, and the latter usually has a pair of yellowish lines
extending from its apex to its posterior ventral angle. The measure-
ments in millimeters of the largest male and largest female are re-
spectively: carapace length 68.0, 81.3; carapace width 46.2, 55.5;
plastron length 46.6, 59.1; depth 29.5, 36.8; length of anterior lobe
of plastron 15.9, 18.9; length of median lobe of plastron 11.9, 16.4;
length of posterior lobe of plastron 19.1, 23.3; width of posterior lobe
of plastron 20.6, 23.3; length of bridge 8.0, 11.5; width of head 16.0,
18.1. The smallest juvenile has a carapace length of 39.0 mm., plas-
tron length of 25.7 mm., depth of 21.0 mm., and head width of 9.5
mm. Because of the paucity of specimens from southern Florida,
comparisons have not been made with northern individuals.
Sternotherus has been taken only in the canals in southern Florida;
the few specimens available have been secured by dipping water hya-
cinths and other aquatic plants from standing water. Although it is
probably more common than the few records indicate, the present
evidence points to the fact that S. odoratus is the least abundant of
the three southern Florida kinosternids. The probable presence of
this turtle in cypress ponds needs verification. Traps baited with
flesh have not succeeded in taking this species in southern Florida.
Sternotherus odoratus occurs in aquatic habitats throughout the
area and has been encountered most frequently in the Everglades;
it does not occur on the Florida Keys. Locality records for 10 speci-
mens are:


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 261

Collier County: 8 mi. WNW Collier-Dade Co. line, Tamiami Trail (1).
Dade County: 23.1 mi. W Miami (2), 24.2 mi. W Miami (1), 33 mi. W Miami (1),
Miami Springs (1), Paradise Key (2). Monroe County: Pinecrest (2).

Kinosternon bauri bauri Garman
Cinosternon baurii Garman, 1891, Bull. Essex Inst., vol. 23, p. 141.
Kinosternon bauri bauri, Uzzell and Schwartz, 1955, p. 33.
The subspecies of Kinosternon bauri have been redefined by Uzzell
and Schwartz (1955) who have shown that the nominal form is re-
stricted to the lower keys. This population is characterized by a
dark carapace, usually having the light lines obscured and the lower
beak either unstreaked or with very weak streaks. Comparisons of
measurements and proportions of specimens from the lower keys
and from the mainland are given in table 9. The largest male of
K. b. bauri is from Stock Island and has a carapace length of 103.9 mm.
Specimens on Big Pine Key were secured by baiting traps with
small mammal carcasses. The turtles came to the bait during day-
light hours, indicating that they may do some of their foraging by
day. Individuals on the lower keys live in small permanent and
temporary pools of fresh and brackish water; observers have noted
them in mangrove swamp. The ability of the animal to survive in
salt water probably has aided it in crossing the narrower channels
between the keys; the distinct markings and coloration of the popu-
lations on the lower keys also tend to demonstrate that these popula-
tions are effectively isolated from populations on the upper keys by
the wide channel today spanned by the Overseas Highway Seven-
Mile Bridge.
As indicated by Uzzell and Schwartz (1955), Kinosternon bauri
occurs only on the oilitic lower keys. Locality records for 33 speci-
mens are:
Monroe County: Big Pine Key (17), Key West (11), 30 mi. N Key West (1),
Stock Island (4).

Kinosternon bauri palmarum Stejneger
Kinosternon bauri palmarum Stejneger, 1925, Jour. Washington Acad. Sci., vol.
15, p. 463. Uzzell and Schwartz, 1955, p. 34.
The mainland and upper key populations of Kinosternon bauri
recognized as the subspecies palmarum by Uzzell and Schwartz
(1955) are characterized by a variably pigmented carapace and a
heavily streaked mandibular beak. The scutes of the carapace are
often transparent, the light color making the light lines on the cara-


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262 BULLETIN FLORIDA STATE MUSEUM Vol. 3

pace indistinct; otherwise the lines are distinct. Measurements and
proportions are given in table 9.

TABLE 9
MEANS AND OBSERVED RANGES OF MEASUREMENTS AND PROPORTIONS OF THE LARGEST
MALES AND FEMALES OF Kinosternon bauri AVAILABLE FROM SOUTHERN FLORIDA.
(15 MALES AND 15 FEMALES FROM THE MAINLAND, 6 MALES AND 15 FEMALES
FROM THE LOWER KEYS).

Character Sex Mainland Lower Keys

Carapace length $ 87.6 ( 80.8-98.7) 89.1 (78.3-103.9)
9 107.6 (101.6-119.0) 89.8 (74.1-110.7)
Carapace width a 56.8 ( 50.4-63.7) 56.9 (50.3-64.6)
9 71.3 ( 68.5-78.3) 59.2 (49.5-73.2)
Plastron length & 70.7 ( 63.3-76.9) 75.6 (68.7-85.7)
9 99.2 ( 92.0-108.8) 79.6 (62.3-99.0)
Greatest depth $ 34.5 ( 30.8-40.1) 33.4 (29.7-38.1)
9 46.3 (41.2-51.3) 36.3 (27.8-48.4)
Length of anterior lobe S 25.4 ( 21.2-28.2) 25.5 (21.8-30.1)
9 31.9 ( 29.6-34.7) 26.2 (21.3-31.7)
Length of median lobe & 20.0 ( 16.8-22.0) 21.1 (19.2-24.0)
9 28.7 ( 25.0-33.7) 22.1 (16.3-28.9)
Length of posterior lobe & 25.4 ( 21.2-28.2) 26.6 (24.3-31.6)
9 37.7 ( 31.8-40.7) 31.2 (24.7-40.1)
Width of posterior lobe & 30.5 ( 24.8-33.8) 32.9 (27.6-38.7)
9 45.0 ( 40.4-47.4) 36.1 (28.8-45.0)
Length of bridge & 14.1 ( 12.4-16.9) 14.9 (12.2-18.4)
9 22.6 ( 19.4-25.2) 16.5 (12.7-23.5)
Length of bridge/ & .160 ( .150-.179) .167 ( .151-.177)
Carapace length 9 .210 ( .190-.231) .183 ( .162-.212)
Length of anterior lobe/ $ 1.76 ( 1.56-1.91) 1.72 ( 1.63-1.86)
Length of bridge 9 1.42 ( 1.26-1.58) 1.61 ( 1.35-2.04)
Carapace width/ & 2.31 ( 2.16-2.42) 2.24 ( 2.07-2.40)
Length of anterior lobe 9 2.24 ( 2.04-2.49) 2.26 ( 2.07-2.38)



Carr (1952: 93) stated that K. bauri "is probably the least aquatic
of the North American kinosternids . It likes to prowl on land
during rains and may frequently be seen in temporary rain pools
after thundershowers." Our observations substantiate this statement;
most of the large series from southern Florida have been taken on
roads and canal banks. Although it may be collected in this matter
at almost any hour, most terrestrial movement seems to occur just at
dusk. Some individuals were collected by dipping water hyacinths
and other aquatic vegetation.


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 263

Kinosternon bauri palmarum is widespread in southern Florida,
occurring in canals, ditches, sloughs, ponds, and larger bodies of
fresh water on the mainland. On the upper keys it sometimes has
been found in brackish water. Locality records for 155 specimens
are:
Broward County: 1 mi. N Dade Co. line (1), 21.5 mi. W Ft. Lauderdale (1),
4.6 mi. S junction Ft. Lauderdale and Okeechobee-Miami roads (1), South New
River Canal at Okeechobee-Miami Road (1). Collier County: 4-7 mi. W Carnes-
town (7), 10.8 mi. W Carnestown (1), Everglades (1), 2 mi. N Everglades (1), 8.2
mi. W Monroe Station (1), 12.6 mi. E Monroe Station (1), 19.6 mi. E Monroe
Station (1), 1 mi. E Naples (1), Ochopee (8), 2.7 mi. W Ochopee (1), 8.2 mi. W
Ochopee (1), 2.7 mi. ESE Royal Palm Hammock (1), 6-9 mi. ESE Royal Palm
Hammock (3), 11-13.5 mi. ESE Royal Palm Hammock (3), Thickahatchee Swamp
(1). Dade County: Biscayne Key (1), Coconut Grove (1), 4 mi. W Florida City
(1), 4.2 mi. S Florida City (22), Homestead (4), Lee Hammock (1), Long Pine Key
(4), 1 mi. N Medley (1), Miami (8), 7 mi. W, 2.5 mi. N Miami (2), 12 mi. W
Miami (3), 13.5-17.5 mi. W Miami (3), 19 mi. W Miami (3), 19 mi. W, 3-6 mi. S
Miami (2), 20-25 mi. W Miami (15), 27-30 mi. W Miami (7), 34-40 mi. W Miami
(7), 2 mi. NW Miami (2), Paradise Key (8), 0.7 mi. E Paradise Key (3), 2.1 mi. E
Paradise Key (1), 1.5 mi. SW Paradise Key (1), 5 mi. SW Paradise Key (1), 10 mi.
SW Paradise Key (1), 17.4 mi. SW Paradise Key (1), Sweetwater (1). Monroe
County: Key Vaca (1), Lower Matecumbe Key (3), Pinecrest (10), Upper Mat-
ecumbe Key (1).

Kinosternon subrubrum steindachneri Siebenrock

The carapace often has three faintly delineated pale longitudinal
stripes, which are not so well developed as the similar stripes in K.
bauri. The plastron may be uniformly black, uniformly yellow, or
any intermediate condition between these extremes; most often it is
horn-colored, with the newer growth showing as dark brown concen-
tric rings surrounding the older, worn, yellowish or horn-colored cen-
tral portions of each lamina. The head is mottled black and whitish;
the upper beak is vertically striped whereas the lower beak is hori-
zontally striped. Both sexes are approximately the same size.
The observed ranges and averages of measurements of five males
and four females (all adult or subadult) from southern Florida are:
carapace length 83.3 to 108.7 (92.8) mm., carapace width 56.2 to 73.8
(63.2) mm., plastral length 67.4 to 88.4 (75.3) mm., greatest depth 34.9
to 45.9 (38.7) mm. Examination of table 10 indicates that in the
three ratios computed there is a tendency for specimens from southern
Florida to average less in the ratio of width of bridge to carapace
length, and to average more in the ratio of anterior lobe of plastron
length to width of bridge. The material from southern Florida more
closely resembles that from northern Florida in all ratios and most


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measurements. In addition, all specimens from peninsular Florida
are distinguishable from specimens from South Carolina on the basis
of the shape of the posterior lobe of the plastron; the Florida speci-
mens have a more or less triangular lobe (apex directed posteriorly),
whereas in South Carolina specimens the posterior lobe is more
quadrate or rectangular. In Florida specimens the greatest width
of the posterior lobe occurs immediately posterior to the posterior
lobe-median lobe suture; in Carolina specimens the greatest width
lies near the midpoint of the free edge of the femoral laminae. Al-
though the comparisons of northern and southern Florida specimens
are based on inadequate samples for statistical analysis, in general
the southern specimens seem to be somewhat smaller.

TABLE 10
MEANS OF MEASUREMENTS AND PROPORTIONS OF Kinosternon subrubrum
FROM THREE REGIONS.

Southern Florida Northern Florida South Carolina

Males 5 2 2
Females 4 4 6
Carapace length 92.8 94.7 80.7
Carapace width 63.2 65.3 55.0
Plastron length 75.3 76.3 71.5
Greatest depth 38.7 40.3 34.8
Length of anterior lobe 27.9 30.5 27.7
Length of median lobe 20.1 20.2 17.5
Length of posterior lobe 27.4 29.7 26.0
Width of posterior lobe 32.3 35.1 33.7
Length of bridge 11.7 12.3 13.6
Width of head 24.9 23.4 18.8
Length of bridge/ .117 .129 .169
Carapace length
Length of anterior lobe/ 2.63 2.42 2.06
Length of bridge
Carapace width/ 2.26 2.28 1.97
Length of anterior lobe


This species is not so abundant in southern Florida as K. bauri;
the two species occur in the same situations. Both have been col-
lected crossing the Tamiami Trail, and all of our specimens from
southern Florida have been taken in this manner. Carr (1940: 99)
stated that the habitat of this turtle is "small streams, sloughs, and
drainage ditches; marshes . ."; we can add little to this since all


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 265


southern Florida specimens have been taken in the immediate vicinity
of canals and other standing water.
Kinosternon subrubrum occurs throughout most of southern Flor-
ida; apparently it avoids the main part of the Everglades, an area
where K. bauri reaches its greatest abundance. This species has not
been recorded from the Florida Keys. A specimen was secured on
Marco Island (which is completely separated from the mainland by
salt water), and its occurrence there may indicate that K. subrubrum
can survive in salt or brackish situations, although it does not usually
occur in saline waters. There is a specimen (CM 26725) from Stock
Island, near Key West, which has been identified as K. subrubrum.
This individual cannot be located at this time, so that the identifica-
tion cannot be verified. It may represent K. subrubrum (and if so,
is the only known record of the species from the Florida Keys), but
it may be assignable to K. bauri, which occurs on Stock Island and
other Florida Keys. Locality records for 19 specimens are:
Collier County: Marco Island (1), Monroe Station (2), 7-10 mi. W Monroe
Station (3), Naples (2), 7.4 mi. SE Naples (2), 5-6 mi. E Ochopee (4), Thicka-
hatchee Swamp (1). Dade County: Miami (2), 30 mi. W Miami (1). Monroe
County: Pinecrest (1).

Terrapene carolina bauri Taylor
Eight males exhibit the following ranges and means of measure-
ments: carapace length 128 to 154 (140.1) mm., carapace width 85
to 104 (95.6) mm., greatest depth 68 to 78 (72.3) mm. Nine females:
carapace length 109 to 142 (127.4) mm., carapace width 74 to 103
(88.7) mm., greatest depth 56 to 76 (68.1) mm. Three males from
the keys have a flare on the posterior half of the carapace; also the
carapace is proportionately wider than in mainland specimens. The
carapace width/carapace length ratio for three males from the keys
is 72.5; that for five males from the mainland is 65.6, and that for
nine females from the mainland is 69.6. There are no females avail-
able from the keys. These data show that in the relatively wider
carapace and presence of a flare posteriorly, the specimens from the
keys resemble Terrapene c. major, thus supporting the evidence given
by Auffenberg (1958). The smallest juvenile examined has a carapace
length of 50 mm., a carapace width of 40 mm., and a greatest depth
of 26 mm.
The series from southern Florida contains many immature indi-
viduals ranging in size from 70 to 100 mm. in carapace length. In
comparing these with the large adults it is noted that there appears


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to be a gradual darkening of the head pattern with age. Usually
there are two longitudinal yellowish-brown, orange, or yellow stripes
on top of the head. In a few specimens these are fragmented into
a row of dashes or dots. In older specimens the stripes are darker,
approaching a dull reddish brown or grayish yellow.
Terrapene reaches its greatest abundance in southern Florida in
the rocky pineland between Miami and Florida City and in the region
where the pineland gives way to the Everglades west of Miami. It
does not inhabit the Everglades. On Marco Island and near Naples
it was found in rosemary scrub, where it is associated with Gopherus
polyphemus. The specimen from Key West was picked up in a vacant
lot in 1954. It seems strange that with all of the available herpeto-
logical material from that island that the species had not previously
been taken there. Since box turtles are often kept as pets, this par-
ticular individual may have been brought to the island from the
mainland. Locality records for 69 specimens are:

Collier County: 2 mi. N Carnestown (1), 6.1 mi. S Deep Lake (1), Marco
Island (1), 7.1 mi. N Jerome (1), 1.8 mi. SE, 1.9 mi. S Naples (1), 24 mi. SE
Naples (1), 8.5 mi. NW Royal Palm Hammock (1). Dade County: 4 mi. S Florida
City (2), 1.4 mi. W, 1.3 mi. S Florida City (1), 5 mi. SW Florida City (3), Home-
stead (2), 4.3 mi. W Homestead (1), 5 mi. W Homestead (1), 9 mi. N, 2 mi. W
Homestead (2), 18 mi. N Homestead (1), 3 mi. W Kendall (1), Long Pine Key
(8), Miami (4), 12-14 mi. W Miami (17), 15-19 mi. W Miami (3), Paradise Key (7),
2 mi. E Peters (1), 2.3 mi. N Richmond Air Base (1), South Miami (1), 2.2 mi. W
Sweetwater (1). Monroe County: Big Pine Key (1), Key Largo (1), Key West (1),
Pinecrest (1), Summerland Key (1).

Malaclemys terrapin macrospilota Hay

The geographical variation and definition of the southeastern sub-
species of Malaclemys terrapin were presented by Schwartz (1955).
Since his discussion no new material has become available; there
are so few specimens of these highly variably turtles from southern
Florida that the delimitation of ranges and the determination of vari-
ation is not possible at this time. Specimens of M. t. macrospilota
from southern Florida are characterized by a dark carapace with
distinct light centers in the central and lateral lamina. No males are
known from southern Florida; four adult females range from 170 to
193 mm. in carapace length.
At Marco Island these turtles have been observed basking on
logs during the day, and as many as six have been seen swimming in
the mangrove-bordered canal. Johnson (1952) reported taking an


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individual on the beach at Key Island, south of Naples, Collier County.
During the period of deposition of eggs (June and July) females oc-
casionally are encountered on roadways and canal banks; other than
this periodic wandering, these turtles are confirmedly aquatic.
In southern Florida M. t. macrospilota is known only from the
mangrove swamps and brackish canals of the Gulf coast as far south
as Marco Island. One specimen of macrospilota purportedly from
Key West (USNM 37021) is of questionable provenance (see Schwartz,
1955: 160). Locality records for 5 specimns are:
Collier County: Marco Island (4), 3.3 mi. SW Royal Palm Hammock (1).

Malaclemys terrapin rhizophorarum Fowler

This subspecies of M. terrapin is distinguished by the presence
of black on the seams of the ventral surface of the marginals at the
level of the bridge, no smudge on the marginals at the bridge, and
head spots fused to form blotches. From the limited number of
specimens available it appears that males are distinctly smaller than
the females; the only male specimen has a carapace length of 117
mm., whereas a female from the same locality (Marquesas) has a car-
apace length of 172 mm.
Our attempts to collect specimens by trapping in mangrove
swamps were futile, as were attempts to secure specimens from fish-
ermen at Key West. Consequently, we have not encountered this
form in southern Florida. Locality records indicate that the sub-
species probably ranges throughout the chain of Florida Keys and
possibly into the region of Florida Bay and the Ten Thousand Islands,
where it may intergrade with M. t. macrospilota. Locality records
for 3 specimens are:
Monroe County: Key Largo (1), Marquesas (2).

Malaclemys terrapin tequesta Schwartz
Malaclemys terrapin tequesta Schwartz, 1955, p. 158.
This subspecies has been adequately discussed and diagnosed by
Schwartz (1955). No further specimens are available from southern
Florida, so that the only known specimen from the area under con-
sideration is the type. To define the southern range of this form ad-
ditional material is needed from the lower reaches of Biscayne Bay
and the extreme northern part of the Florida Keys. The single avail-
able specimen is from Miami Beach, Dade County.


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Pseudemys floridana peninsularis Carr
Two adult females from Dade and Broward counties have the
following measurements (in millimeters): carapace length 287 and
320, carapace width 205 and 222, plastron length 262 and 290, depth
of shell 127 and 141, width of head 36 and 38, length of bridge 111
and 118; a small male from Collier County has these: carapace length
206, carapace width 147, plastron length 176, depth of shell 81, width
of head 27, length of bridge 60.
Usually the carapace is dark brown to black with a complex yel-
low reticular pattern. The tomium of the upper jaw is noncusped
and nonserrate, thus having a smooth appearance. This character
will distinguish the species from P. nelsoni, which usually has a ser-
rate and cusped upper tomium.
Pseudemys floridana is seldom encountered with the frequency of
P. nelsoni, but certainly the former is more abundant than the few
specimens indicate. They inhabit the canals and sloughs of the
Everglades, but have not been observed in cypress ponds. Most
frequently they are seen basking with P. nelsoni on mats of vegeta-
tion or rocks and partially submerged logs and branches. Both spe-
cies are equally shy and cannot be approached.
The subspecies P. f. peninsularis occurs throughout the Ever-
glades and adjoining fresh waters of southern Florida; there are no
specimens available to us from the keys, although Carr (1935) re-
ported the occurrence of this turtle on Key Largo. Locality records
for 11 specimens are:
Broward County: No additional data (1). Collier County: 10 mi. N Cope-
land (1), Royal Palm Hammock (1). Dade County: No additional data (4), 7 mi. W
Medley (1), Paradise Key (3).

Pseudemys nelsoni Carr
Females attain a greater size than males; the following measure-
ments of 11 adult females from southern Florida show the large size
and variation: carapace length 251 to 320 (284) mm., carapace width
176 to 216 (198) mm., plastron length 234 to 294 (265) mm., depth
of shell 108 to 139 (126) mm., length of bridge 96 to 124 (108) mm.,
width of head 34 to 42 (37) mm. Measurements, in millimeters, of
two males from Dade County are: carapace length 229 and 170,
carapace width 163 and 133, plastron length 209 and 163, depth of
shell 94 and 76, length of bridge 81 and 66, width of head 31 and 26;
a juvenile with a prominent umbilical scar from Collier County has


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 269

these: carapace length 32.4, carapace width 29.6, plastron length
28.9, depth of shell 16.8, length of bridge 9.7, width of head 9.0.
There is at hand an adult female turtle (ChM 54.144.19) which has
been pronounced by John W. Crenshaw, Jr. a hybrid between P. nel-
soni and P. floridana peninsularis. This specimen has a carapace
length of 292 mm., and was taken 23 June 1954, 7 miles west of Med-
ley, Dade County. This is one individual from a series of P. nelsoni
and P. floridana collected at this locality by overtaking the turtles as
they basked on a mud dam, before they could return to the water
and escape. Twelve eggs were removed from this female in late
June; they hatched on 10 September. This lot of 12 hatchlings has
been studied by Crenshaw who concluded that they show the hybrid
characteristics of the adult female.
Pseudemys nelsoni is a confirmed basker; large numbers can be
seen during the summer on logs and rocks in the Tamiami Canal.
That such basking activity is not confined to warmer months is dem-
onstrated by basking as well as foraging individuals being seen in
late December at Paradise Key. This species is an inhabitant of
the canals and sloughs of southern Florida; it does not occur on the
keys. Locality records for 40 specimens are:
Collier County: 5 mi. SE Naples (1). Dade County: 10.1 mi. NE Coot Bay
Ranger Station (1), 7 mi. W Medley (9 nelsoni, 13 floridana X nelsoni), Miami
(5), 11 mi. W Miami (1), 18-23 mi. W Miami (4), 30 mi. W Miami (1), 36 mi. W
Miami (3), 2.6 mi. SW Paradise Key (1). Monroe County: 1.2 mi. E Pinecrest (1).

Deirochelys reticularia chrysea Schwartz
Deirochelys reticularia chrysea Schwartz, 1956a, p. 476.
Measurements for four males from southern Florida are: carapace
length 110.2 to 144.0 (124.0) mm., carapace width 82.6 to 99.4 (85.8)
mm., greatest depth 45.9 to 56.0 (49.7) mm. The same measurements
for the single female examined are: 187, 126, and 82 mm. The vari-
ation in coloration in Florida and the differences between Floridian
specimens and those from other parts of the range have been dis-
cussed at length by Schwartz (1956a).
Deirochelys is generally considered to be an inhabitant of ponds,
and our meager data from southern Florida indicate that in this area
it inhabits standing water or canals with a low gradient. It has been
observed crossing the Tamiami Trail and occasionally basking on
logs in the Tamiami Canal. Trapping in canals in southern Florida
has yielded no specimens. Elsewhere it has been taken in cypress
ponds, and probably it does not shun this habitat in southern Florida.


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A female (185 mm. in carapace length) was encountered on 15
November 1952, near Monroe Station on the Tamiami Trail, at 1:30
p.m., while depositing eggs. The hole for egg deposition was cylin-
drical, approximately four inches deep and three inches in diameter
at its mouth. One egg had been deposited at the time of the dis-
covery of the turtle, and three additional ones were played in cap-
tivity. The four eggs averaged 33.1 mm. in length, with an observed
range of 32.0 to 34.1 mm. These are smaller than those reported by
Carr (1952: 319), who stated that the eggs of this species vary between
37 and 40 mm. Carr likewise reported that the number of eggs per
clutch may vary from 7 to 15, and that in Florida this species may
deposit its eggs anytime of the year.
Deirochelys appears to be rare in southern Florida; doubtless its
rarity is more apparent than real. It probably ranges throughout the
Everglades; it does not occur on the keys. Locality records for 7
specimens are:

Broward County: New River (1). Collier County: 2 mi. N Copeland (1), 5.8
mi. E Monroe Station (1). Dade County: 5 mi. W Florida City (1), Long Pine
Key (1), Miami (1), Tamiami Trail (1).

Gopherus polyphemus Daudin

Gopherus apparently is a rare turtle in southern Florida. A single
adult female from the vicinity of Naples, Collier County, has a car-
apace length of 206 mm., carapace width 149 mm., greatest depth 88
mm., and plastron length (along midline) 185 mm.
In southern Florida the gopher turtle is confined to the sandy
scrub areas that barely enter southern Florida in scattered patches in
Broward and Dade counties on the east and along the Gulf coast as
far south as Marco Island on the west. Although no specimens were
secured, burrows belonging to this turtle were observed on Marco
Island. Locality records for 3 specimens are:

Broward County: Hollywood (1). Collier County: 7.4 mi. N Naples (1).
Dade County: Miami (1).

Chelonia mydas Linnaeus

The sea turtles have not been studied by us; however, we have
attempted to compile locality records for existing specimens from
southern Florida. These are included solely for the sake of complete-
ness of the herpetological record for southern Florida. Five speci-
mens of this species are available from Key West, Monroe County.


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Eretmochelys imbricata Linnaeus
Locality records for 9 specimens are:
Southern Florida (1). Monroe County: Garden Key (Tortugas) (1), Key
West (7).
Lepidochelys olivacea kempi Garman
One specimen is available from Sands Key, Dade County, and
one from Key West, Monroe County.

Caretta caretta Linnaeus
Locality records for 23 specimens are:
Dade County: Governor Pass near Miami (1). Monroe County: "Florida
Keys" (1), Key West (6), Long Key (2), Tortugas (13).

Trionyx ferox Schneider
Testudo ferox Schneider, 1783, Naturgeschichte Schildkr6ten, p. 330. Schwartz,
1956b, p. 1.
In adults the carapace is dull brown or blackish, and the skin is
likewise dark in color. In juveniles, however, the dorsal pattern is
made up of a series of bluish-black blotches on a dull brown back-
ground. The carapace is edged with orange, grading to yellow pos-
teriorly. The plastron and ventral surface of the carapace are uni-
formly slate gray. Measurements of the largest female and the only
male studied are: carapace length 321 and 183 mm., carapace width
231 and 140 mm., plastron length 194 and 116 mm., greatest depth
82 and 49 mm., length of bridge 38 and 20 mm., width of head 46
and 30 mm.
When specimens from southern Florida are compared with series
from throughout the range of the species (Georgia, South Carolina,
and northern and western Florida), little difference is observed be-
tween southern specimens and those from the rest of the range (see
Schwartz, 1956b). Crenshaw and Hopkins (1955: 19) have noted that
specimens of T. ferox from southern Florida (Lake Okeechobee south-
ward) have a greater carapace width relative to head width. Color
pattern and proportions are somewhat similar when specimens from
southern Florida and South Carolina are compared. i
Occasional individuals are observed basking. Often in preference
to climbing on banks these turtles rest on beds of aquatic vegetation.
They are always wary and quickly dive when approached. Adults
and subadults are sometimes encountered on roads bordering canals;


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adult females leave the water to deposit eggs and thus are readily col-
lected. Although these turtles are omnivorous, traps baited with fish
and beef have not taken them. They are often taken by fishermen
on hooks baited with worms.
Trionyx occupies all fresh water habitats and has been noted in
brackish water (Carr, 1952: 417) and even in marine situations (Carr,
1952; Neill, 1951a: 16). Although we have not taken specimens from
cypress ponds, these turtles undoubtedly occur in that habitat. They
are abundant in the Tamiami Canal and other artificial fresh-water
canals. They are absent from the Florida Keys. Locality records
for 15 specimens are:
Broward County: Birch State Park (1), Ft. Lauderdale (1), 22 mi. WNW, 6
mi. SSE Ft. Lauderdale (1). Collier County: 11.2 mi. E Monroe Station (1). Dade
County: Miami (1), 15 mi. NW Miami (2), 19 mi. W Miami (2), 35 mi. W Miami
(1), 42 mi. W Miami (1), 40 mi. W, 1.6 mi. NW Miami (1), Paradise Key (2),
West Miami (1).

Dermochelys coriacea Linnaeus
Three specimens are available from Dade County-two from Bis-
cayne Bay, one from Miami Beach.

Crocodylus acutus Cuvier
Moore (1953) has thoroughly reviewed the status of the crocodile
in southern Florida. He has given the most comprehensive data on
habits and life history of the species in Florida yet presented. Moore's
data are in conflict with certain information presented by Carr (1940:
68-9). Carr stated that Crocodylus lives in fresh and salt water; ap-
parently the species (as it now exists) is almost entirely restricted
to brackish and salt-water habitats. We have observed it in brackish
West Lake near Coot Bay Ranger Station in the Everglades National
Park. Moore thinks that the abundance of Alligator in the fresh-
water habitats may have restricted Crocodylus to those of salt water.
Dates for egg laying in captivity range from 12 March to 10 May;
21 to 56 eggs may be laid (Moore, 1953). He described the nest as
a pile of sand lacking vegetation. Carr (1940) reported 15 to 20
young and an adult in a drainage ditch on Lower Matecumbe Key.
Although Carr recorded Crocodylus from Collier County, we have
been unable to find any specimens from that area. Moore stated
that he did not believe that the crocodile occurred on the lower west
coast of Florida. Apparently the range is restricted to the brackish
and marine habitats from Cape Sable eastward along the southern


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 273

coast of the peninsula, across Florida Bay and as far south as Big
Pine Key. The Big Pine Key specimen was collected in 1953. The
range, within recent years, extended northward along the east coast
into Biscayne Bay. For locality records other than those given below,
see Moore (1953). Locality records for 12 specimens are:
Dade County: Biscayne Bay (3), "Everglades" (5). Monroe County: Big Pine
Key (1), Cape Sable (2), Key Largo (1).

Alligator mississipiensis Daudin
Our experience with this species in southern Florida has been
limited to observations and sight records; none was collected. Since
the alligator has been protected it seems to be increasing in numbers
in the Everglades, and especially in the Everglades National Park.
Individuals often find their way into small ponds in parks in Miami
and Coral Gables. Numerous individuals have been observed from
the road from Paradise Key to Flamingo, from the Loop Road, and
from the Tamiami Trail. In the slough at Paradise Key as many as
nine individuals have been seen at one time. In the sloughs and canals
in the Everglades the alligator inhabits fresh water. Often small in-
dividuals are seen resting on mats of aquatic vegetation. Several
individuals have been observed in the mangrove-bordered, brackish
West Lake near Coot Bay Ranger Station in the Everglades National
Park. Here they occur in the same habitat with Crocodylus.
The largest specimen observed was approximately 10 feet in total
length; doubtlessly larger individuals exist in the more inaccessible
regions of the Everglades.
Aside from the few locality records given below that are based
on museum specimens, the many reports of this species in the litera-
ture show that it has a more extensive range in southern Florida.
Fowler (1906) reported it from Little Pine and Summerland keys;
Small (1923: 212) observed the species on Big Pine Key. Allen and
Slatten (1945) reported the capture of three specimens at night in salt
water at Key West. Locality records for 17 specimens are:
Collier County: 10 mi. N Everglades (2). Dade County: near Homestead
(13), Monroe County (2).

Gonatodes fuscus Hallowell
Introduced into Key West sometime prior to April 1939, when the
first specimens were collected (Carr, 1939), a colony of these geckos
has thrived on the island ever since. In 1939 the species was abundant


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in the old freight yards; today they are still found there, less commonly
in the Navy Yards and in downtown Key West. The fact that numer-
ous young individuals are seen indicates that the population is thriving.
The largest male examined has a snout-vent length of 40.3 mm.
and an incomplete tail; another with a snout-vent length of 38.5
mm. has a tail length of 46.6 mm. The largest female has a snout-
vent length of 37.8 mm. and a tail length of 30.5 mm.
Gonatodes fuscus is restricted to edificarian situations on Key
West. Because of its habits it may be transported in the future to
other cities, but it is unlikely that it will become established in natural
habitats in southern Florida. Eleven specimens are available from
Key West, Monroe County.

Hemidactylus turcicus turcicus Linnaeus
Fowler (1915: 252) reported Hemidactylus mabouia from Key West.
This specimen, collected in March 1910, cannot be located to check
its identity. Probably the specimen is an example of Hemidactylus
turcicus, and, if so, stands as the first record for the species from Key
West. Stejneger (1922) reported H. turcicus from Key West, the speci-
mens having been collected in 1915 and 1921, and Barbour (1936;
113) recorded the species for the first time from Miami. We have
found Hemidactylus to be abundant in Key West, but notably less
so in Miami, there having been only five specimens collected in
Miami in recent years. In the small cemetery in the middle of the
city of Key West these lizards abound on stone walls, tombstones,
and large urns. Numerous young have been collected, and clutches
of eggs have been found in the bottoms of the urns.
The largest male has a snout-vent length of 55 mm., the largest
female 53 mm. A juvenile has a snout-vent length of 23.6 mm.
From our experience with this species in Miami and Key West it
is confined to edificarian situations. Two specimens (USNM 101148-
149) from Big Pine Key may have been collected in the pine forest on
that island. We have been unsuccessful in discovering this lizard in
natural areas anywhere in southern Florida. Locality records for 88
specimens are:
Dade County: Miami (5). Monroe County: Big Pine Key (2), Key West (81).

Sphaerodactylus argus argus Gosse
Sphaerodactylus argus Gosse, 1850, Ann. Mag. Nat. Hist., ser. 6, vol. 2, p. 347.
Sphaerodactylus argus argus, Savage, 1954, p. 327.


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1958 DUELLMAN AND SCHWARTZ: AMPHIBIANS AND REPTILES 275

A native of Jamaica, this species apparently was introduced into
Cuba and Key West (Savage, 1954: 328). L. Neil Bell stated (in litt.)
that of more than 150 specimens of Sphaerodactylus from Key West
examined by him, none was identified as S. argus. The three speci-
mens reported on by Savage (SU 10439-40, 10442) were collected in
Key West by J. L. Grimmer and Earl S. Herald in February 1944.
Recent collecting has shown no evidence of this species, and we as-
sume that although it may have been introduced and established for
a short period of time, no colony of Sphaerodactylus argus is extant
on the island now.

Sphaerodactylus cinereus Wagler
Stejneger (1922) first reported this lizard from Key West, one
specimen having been collected there in 1921. Burt (1937: 352) re-
corded a specimen from Key Largo, collected in 1928. This specimen
(USNM 75113), although dried, can be identified as Sphaerodactylus
notatus. The main colony of these lizards is in Key West; however,
in 1954, a single specimen was secured on Boca Chica Key, two islands
removed from Key West and connected with it by a causeway.
On Key West the lizards are found commonly in the old cemetery
and on buildings in town. They are secretive and wary, and because
of their small size are difficult to capture. The largest male has a
snout-vent length of 33.5 mm., the largest female, 35 mm. Numerous
brightly banded juveniles with brick red tails were observed in the
cemetery. The smallest one collected has a snout-vent length of
16.5 mm.
Like the preceding species of geckos, all introduced into southern
Florida in relatively recent years, S. cinereus is an inhabitant of build-
ings in cities; none has been found in natural areas. Of 78 available
specimens, one is from Boca Chica Key, and the remainder from Key
West, Monroe County.

Sphaerodactylus notatus Baird
The pigmentation of this small gecko is quite variable, and there
is a distinct difference in coloration between the sexes. In males the
entire dorsal surface of the body is spotted with dark punctations,
each of which consists of a single darkened scale. There is no tend-
ency for the head spots to be arranged in either transverse or longi-
tudinal rows or to form stripes. The gular region is strongly pig-
mented with dark spots. The preanal patch of thickened scales is


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well defined. Females have three longitudinal dark stripes on the
head, one median and two postocular. The centers of these stripes
are lighter than the borders; this is especially true of the median
stripe. Posterior to the scapular region the dorsum has the same
dark punctations as in the males. Smith (1946: 75) mentioned speci-
mens that were intermediate between these patterns as well as uni-
color individuals. The latter are old males in which all traces of
the spots have been lost. By utilizing the large series of available
specimens, so-called intermediate specimens may be placed with
ease. Males always have spotted heads; in females, the dark lines
or rows of spots or dashes, perhaps ill-defined, are always present.
The largest specimen examined had a total length of 30.5 mm.;
the smallest sexually mature individual was 20.5 mm. long.
Sphaerodactylus notatus has been collected in pineland, hammocks,
cypress heads, vacant city lots, in deserted houses, and in inhabited
wooden buildings. It is abundant at Matheson Hammock, Paradise
Key, Key Largo, and Key West. Apparently the species is primarily
diurnal, although occasional individuals have been observed at night.
The breeding habits of S. notatus have received little mention in
the literature. Mittleman (1950) reported an egg in a specimen pre-
sumably collected on 6 May 1948. The egg measured 3 by 5.5 mm.
Carr (1940: 71) stated: "The eggs are laid from June to August,
singularly or by twos or threes . ." On 29 December 1951, seven
S. notatus eggs were found at Key West; the average size was 4 by
6 mm. Five of these eggs hatched in 74 days, the other two required
79 days. On 8 March 1952, a large composite nest containing 280
eggs was found in a termite-ridden log at Key West. These eggs
had an average size of 4.3 by 5.8 mm. and hatched over a period of
two months. The eggs are hard, oval, and with a brittle shell. Dissec-
tion of gravid females indicates that only one egg matures at a time.
It is probable that a single female deposits several eggs each year,
but whether breeding occurs more than once a year is problematical.
Egg laying, as demonstrated by the large number found in one log,
is often communal; a single egg is rarely found. Eggs have been
found in a variety of sites, including bases of palm fronds, termite-
ridden log, driftwood, leaf debris, rotting logs, beneath stones, in piles
of boards, coconut husks, walls of a deserted cistern, and beneath tar
paper on roofs of houses.
Examination of stomach contents has revealed the following food
items: Coleoptera (Carabidae, Chrysomelidae, Curculionidae, Hydro-
philidae, Scarabaeidae, Scolytidae, Staphylinidae), Hymenoptera (For-
micidae), Hemiptera, lepidopteran larvae, and one annelid.


Li


Vol. 3




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