THE SYSTEMATICS AND PALEOECOLOGY 7
OF TWO LATE PLEISTOCENE ‘ . V
HERPETOFALFNAS FROM, THE ,-

SOflTHEASTERN UNETED STATES ‘ ' 7

Dissertatian for the ,Degree of Ph‘ D.
MECHIGAN STATE UNEVERSETY
VENCENT VECTOR WILSOfi
197.5

LIBRARY

Michigan State
University

6?; .

 
 
 
 

VFW

This is to certify that the

thesis entitled

The Systematics and Paleoecology of two Late
Pleistocene Herpetofaunas from the Southeastern
United States '

presented by

Vincent Victor Wilson

has been accepted towards fulfillment
of the requirements for

 

 

 

 

PhD degree in Zoology
/ vAMajorKygfe/
. L// J. Alan Holman
Date 17 Aprll, I975
0-7639

 
       

  

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ABSTRACT

THE SYSTEMATICS AND PALEOECOLOGY OF TWO LATE
PLEISTOCENE HERPETOFAUNAS FROM THE
SOUTHEASTERN UNITED STATES

BY

Vincent Victor Wilson

Fossils representing late Pleistocene herpetofaunas
from Reddick, Florida, and Ladds, Georgia, are identified
and reviewed. The known number of reptiles and amphibians
is increased from thirty-nine to forty-four at the Reddick
site, and is increased from twenty-three to twenty-six at
the Ladds site.

Each herpetofauna is discussed separately. Special
attention is given to the paleoecological and paleoclimatic
significance of the herpetofaunas. Evidence is presented
that the winters at Reddick were less cold at the time of
fossil deposition than they are now. Little faunal
evidence was found to indicate that the late Pleistocene
climate at Ladds differed substantially from that of the

present.

THE SYSTEMATICS AND PALEOECOLOGY OF TWO LATE
PLEISTOCENE HERPETOFAUNAS FROM THE

SOUTHEASTERN UNITED STATES

BY

Vincent Victor Wilson

A DISSERTATION

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

DOCTOR OF PHILOSOPHY

Department of Zoology

1975

For Sally

ii

ACKNOWLEDGMENTS

I would like to thank my major professor, J. Alan
Holman, and my doctoral committee, Donald L. Beaver,
Jane E. Elliott, and Marvin M. Hensley, for their help in
the preparation of this paper. I would also like to thank
Ms. Chris Kulczcki who prepared the maps. Appreciation is
also expressed to Lewis Lipps and Clayton Ray of the
National Museum of National History for the loan of the

Ladds fossils.

iii

TABLE OF CONTENTS

Page

LIST OF TABLES . . . . . . . . . . . . . v

LIST OF FIGURES. . . . . . . . . . . . . vi

INTRODUCTION. . . . . . . . . . . . . . 1
PART I. THE PLEISTOCENE HERPETOFAUNA OF REDDICK IB,

FLORIDA I O O O O o O O O O 0 O O O 2

ANNOTATED LIST. . . . . . . . . . . . 5

DISCUSSION . . . . . . . . . . . . . 32

The Herpetofauna. . . . . . . . . . 32

Paleoecology . . . . . . . 33

Zoogeography and Pleistocene Climate . . . 37

SUMMARY 0 o o o o o o o o o o o o o 4 4
PART II. THE PLEISTOCENE HERPETOFAUNA OF THE LADDS,

GEORGIA O O O I O O O O O O I O O O 4 5

ANNOTATED LIST. . . . . . . . . . . . 47

DISCUSSION . . . . . . . . . . . . . 60

The Herpetofauna. . . . . . . . . . 6O

Paleoecology . . . . . . . . . . 6O

Zoogeography and Pleistocene Climate . . . 61

SUMMARY . . . . . . . . . . . . . . 63

LITERATURE CITED . . . . . . . . . . . . 64

iv

LIST OF TABLES

Table Page

1. Habitat preferences of the Reddick lB
herpetofauna . . . . . . . . . . 36

Figure

1.

2.

LIST OF FIGURES

Reddick amphibian fossils . . . . . .
Reddick anuran fossils . . . . . . .
Reddick amphibian and reptile fossils . .
Reddick snake fossils . . . . . . .
Reddick snake fossils . . . . . . .

Southern range limits of Carphophis amoenus

 

and Bufo w. fowleri. . . . . . .
Ladds amphibian and reptile fossils. . .

Range limits of Clemmys insculpta and Bufo
terrestris. . . . . . . . . .

 

 

vi

Page

12
18
26

30

38

51

62

INTRODUCTION

Hibbard (1970) presented an interpretation of the
Pleistocene and recent environments of the central great
plains. His analysis was based primarily on the wealth of
data available on the successive faunas from late Cenozoic
Kansas and.Oklahoma.

The same type of interpretation remains to be done
for the southeastern United States. Unfortunately, the
Pleistocene faunas of this area have not been studied to
the extent of their Great Plains counterparts. This study
makes a small attempt to correct that situation.

In the southeastern United States, only northern
Florida has been found to have an extensive series of
Pleistocene deposits containing local faunas. The largest
known Pleistocene fauna is found near Reddick, Marion
County, Florida. Only one site containing a sizable
Pleistocene terrestrial vertebrate fauna has been reported
from another southeastern state. This is the Ladds, Bartow
County, Georgia deposit. The study of the herpetofaunas of

these two deposits constitutes the basis of this report.

PART I
THE PLEISTOCENE HERPETOFAUNA OF

REDDICK IB, FLORIDA

Reddick I is an abandoned limerock quarry located
one mile southeast of Reddick, Marion County, Florida
(SW 1/4, NW 1/4, Sec. 14, T. 13 S., R. 21 E.). The quarry
is currently on a portion of prOperty owned by Mr. Jimmy
Williams.

As a result of previous quarrying Operations, four
major fossil bearing deposits lie exposed at Reddick I.

Gut and Ray (1963) and Hamon (1964) have designated these
four deposits as IA, IB, IC, and ID. The fossils occur in
the mostly unconsolidated earths which are found in the
solution pipes and caverns of the area.

Hamon (1964) describes Reddick IB as a sink
filling. The fossils found there probably represent
animals that were trapped by falling into, or by taking
refuge in the solution pipe(s) of the area. Although other
vertebrate fossils occur in the residual clay and sand of
the 18 site, they are outnumbered by the high concentration

of reptile and amphibian fossils found there.

A number of workers have investigated various
segments of the Reddick I herpetofauna. Auffenberg (1955,
1956, 1958, 1959, and 1963a and b) has identified the
majority of the reptiles found there. Goin and Auffenberg
(1955) record an amphibian from the 18 site. Holman
(1962a) lists several reptiles and amphibians from the
IC site.

Detailed stratigraphic treatment of the area can be
found in Auffenberg (1963b), Bader (1957), Brodkorb (1957),
and Hamon (1964). Auffenberg (1963b) and Brodkorb (1957)
have indicated that the site is of Illinoin age. Pinkham
(1971), using recent stratigraphic evidence provided by
Brooks (1968) as well as correlation by faunas, concludes
that Reddick is of Sangamon age.

In June and July, 1968, Dr. J. Alan Holman and
Clair Ossian processed matrix and collected fossils from the
IB site. A volume of approximately one cubic yard of matrix
was removed from one exposure of the fossil bearing stratum.
This matrix was processed using a modified version of the
washing and screening method of Hibbard (1949). Thousands i
of vertebrate fossils were recovered by this method and
were eventually sorted into their apprOpriate class. A
list of the amphibians and reptiles found in this sample

follows.

Class Amphibia

 

Order Urodela

Family Sirenidae
Siren lacertina Linnaeus
Family Ambystomatidae
Ambystoma tigrinum (Green)

 

 

Order Anura

Family Pelobatidae

Scaphiopus holbrooki (Harlan)
Family Ranidae

Rana r 110 Stejneger

Rana pip1ens Schreber

Rana aerolata Baird and Girard
Family Microhylidae

Gastrophyrne carolinensis (Holbrook)
Family Bufonidae

Bufo terrestris (Bonnaterre)

Bufo woodhouséi fowleri (Hinckley)

Bufo quercicus (Holbrook)
Family Hylidae

Hyla cinerea (Schneider)

Psuedacris nigrita (Le Conte)

 

 

 

 

 

 

 

 

 

Class Reptilia
Class Reptilia

 

Order Testudines

Family Emydidae
Terrapene carolina (Linnaeus)
Family Testudinidae
Gopherus polyphemus (Daudin)
Geochelone crassiscutata (Leidy)
Geochelone incisa (Hay)

 

 

 

 

Order Squamata

Family Iguanidae

Anolis carolinensis (Voigt)
Family Teidae

Cnemidophorus sexlineatus (Linnaeus)
Family Anguidae

Qphisaurus ventralis (Linnaeus)
Family Scincidae

Eumeces sp. indet.

 

 

 

Family Colubridae
Qarphophis amoenus (Say)
Ela he sp. indet.
Pituophis melanoleucus (Daudin)
Coluber or Masticophis
Drymarchon corais (Daudin)
Heterodon platyrhinos (Latreille
Heterodon Simus (Linnaeus)
Storeria dekayi (Holbrook)

Thamno his sirtalis (Linnaeus)
Famin Elapidae
Micrurus fulvius (Linnaeus)

Famin Viperidae
Sistrurus miliarius (Linnaeus)
Crotalus adamanteus Beauvois

 

 

 

 

 

 

 

 

 

 

In the annotated list that follows, if the Species
under consideration has not been reported from 13 previously,
reference is given to the closest known Pleistocene
occurrence of that Species. This procedure is followed
in all Orders but the Squamata. Since Auffenberg (1963b)
has previously reported all of the snakes from the IB site,

his record is not repeated under each Species account.

ANNOTATED LIST

Class Amphibia

 

Order Anura
Family Sirenidae

Siren lacertina Linnaeus

Materia1--Eleven thoracic verbebrae, MSU 640,
figures 1A, 113, 1c, 10.

Remarks--These typical §i£2n_vertebrae are assigned
to Siren lacertina because of their large size. Vertebrae
of S. intermedia Le Conte are much smaller. In the two

skeletons of S. intermedia available for study, the haemal

 

Figure 1. Reddick amphibian fossils.

A. Fossil Siren lacertina vertebra, anterior, x 2.9.

 

B. Fossil Siren lacertina vertebra, lateral, x 3.3.

 

C. Fossil Siren lacertina vertebra, dorsal, x 3.3.

 

D. Fossil Siren lacertina vertebra, ventral, x 3.3.

 

E. Recent Siren lacertina vertebra, ventral, x 4.7.

 

F. Recent Siren intermedia vertebra, ventral, x 4.7.

 

G. Fossil Ambystoma tigrinum vertebra, lateral, x 5.0.

 

H. Fossil Scaphiopus holbrooki sphenethmoid, x 2.3.

 

I. Fossil ScaphiOpus holbrooki frontoparietal, x 3.2.

 

J. Fossil Scaphiopus holbrooki nasal, x 3.5.

 

K. Fossil Scaphigpus holbrooki ilium, x 3.0.

 

L. Fossil Rana grylio maxilla, x 2.5.

 

 

M. Fossil Rana grylio humerus, x 2.3.

 

 

Figure 1. Reddick amphibian fossils.

keel of the vertebrae are wider and do not extend as close

to the lip of the cotyle as in S. lacterina (Figures 13,

 

1F). The fossil vertebrae resemble S. lacertina in these

 

characters.

Goin and Auffenberg (1955) record S. lacertina

 

from this site.

Family Ambystomatidae
Ambystoma tigrinum (Green)
Material--Thirty-nine vertebrae, MSU 641, figure 16.

Remarks-~The vertebrae of Ambystoma tigrinum are

 

distinct. Besides exhibiting the wide and short prOportions
as defined by Tihen (1958), the posterior portion of the
neural arch is directed upward and extends well beyond the
end of the centrum. Holman (1969) has reported the latter
feature to be characteristic of S. tigrinum.

This is the first report of this Species from this

site.

Order Anura
Family Pelobatidae
chphiopus holbrooki (Harlan)

Materia1--One nasal, seven front0parietals, two
sphenethmoids, three left ilia, six right ilia, one
urostyle, MSU 649, figures 1H, 11, lJ, 1K.

Remarks-~The identified elements are distinct and

not easily confused with other species.

This species was first reported from Reddick 18 by

Gut and Ray (1963).

Family Ranidae

Rana grylio Stejneger

 

Material--One maxilla, one pterygoid, one dentary,
one scapula, one coracoid, two humeri, one radioulna, one
sacrum, two ilia, one femur, one urostyle, MSU 646,
figures 1L, 1M.

Remarks--These fossils apparently represent one
large male. The entire series was recovered from one small
clump of matrix. Species identification was based on
several of the elements. The ilia indicate the 3223

catesbeiana Shaw, 3. hecksheri Wright, S. grylio group of

 

 

Auffenberg (1957). The maxilla indicates grylio. The
section of the maxilla anterior to the nasal process is

pr0portionately higher and shorter in catesbeiana and

 

heckscheri than in grylio. In this character the fossil

 

maxilla resembles 3. grylio.
Sex was determined by the presence of a well

developed epicondylar flange on the humerus. Chantell

(1970) verified the value of this feature in determining
sex.

This is the first fossil record of S. grylio from
Reddick lB. Tihen (1952) records 3. grylio from the

Pleistocene of Alachua County, Florida.

10

Rana pipiens Schreber

 

Material--Eight left ilia, seven right ilia, MSU
644, figure 2B.

Remarks--Auffenberg (1957) divides the ilia of New
World 3222 into two morphological groups based on the slope
of the posterodorsal border of the ilial crest. In the
pipiens group the slope is more gradual than in the

catesbeiana group. The fossils show the pipiens group

 

character. The only two members of the pipiens group
appearing in or near Florida today are S. pipiens and S.
areolata Baird and Girard. S. areolata ilia can be
distinguished from those of pipiens by the more pronounced
crainiad expansion of the ventral acetabular expansion that
occurs in the latter. This character can be seen by com-
paring the reference ilia of S. pipiens and S. areolata
shown in figures 2A and 2C. The fossil ilia resemble
pipiens.

Holman (1962a) records S. pipiens from this site.

Rana areolata Baird and Girard

 

Material--Three left ilia, four right ilia, MSU
645, Figure 2D.

Remarks--As previously discussed under Rana pipiens,

 

figures 2A and_2C illustrate the difference between the

ventral acetabular expansion of Rana pipiens Schreber and

 

S. areolata. This difference was found to be constant in

Figure 2.
A. Recent
B. Fossil
C. Recent
D. Fossil
E. Fossil
F. Fossil
G. Fossil
H. Fossil
I. Fossil
J. Fossil
K. Fossil
L. Fossil
M. Fossil
N. Fossil

Rana

11

Reddick anuran fossils.

pipiens ilium, x 3.0.

 

Rana

pipiens ilium, x 3.0.

 

Rana

areolata ilium, x 3.0.

 

Rana

areolata ilium, x 3.0.

 

GastroPhyrne carolinensis ilium, x 4.0.

 

Bufo

terrestris nasal, x 3.5.

 

Bufo

terrestris squamosal, x 3.5.

 

Bufo

terrestris frontOparietal, x 3.1.

 

Bufo

woodhousei fowleri ilium, x 5.5.

 

Bufo

terrestris ilium, x 5.5.

 

Bufo

quercicus ilium, x 5.0.

 

Bufo

woodhousei fowleri ilium, x 5.5.

 

Bufo

terrestris ilium, x 5.5.

 

Hyla

cinerea ilium, x 4.4.

 

12

 

Figure 2. Reddick anuran fossils.

13‘

the nine skeletons of areolata and seventy skeletons of
pipiens available for study.

This is the first fossil record of S. areolata.

Family Microhylidae

GastroPhryne carolinensis (Holbrook)

 

Material--Ten left ilia, four right ilia, MSU 651,
figure 23.

Remarks--The ilia of GastrOphryne carolinensis are

 

 

easily recognized by their rectangular ventral acetabular
expansion.
This species has been reported from Reddick 13 by

Holman (1962a).

Family Bufonidae

 

Bufo terrestris (Bonnaterre)

Material--One nasal, twenty-three frontOparietals,
seven squamosals, twelve left ilia, fifteen right ilia,
MSU 642, figures 2F, 26, 2H, 2J, 2M.

Remarks-—Tihen (1962) states that in EEEQ
terrestris the height of the ilial prominence is never more
than 30 percent of its base. Measurements made on fourteen
recent MSU S. terrestris skeletons (twenty-eight ilia)
support his findings. The MSU ilia showed a range of
twenty-one to twenty-nine percent for this character. All
fossil 2232 ilia with ilial prominence height to base

ratios of less than 30 percent were assigned to

14

S. terrestris. All other bufonids of the eastern United

 

States show ratios above 30 percent (Tihen, 1962).
The frontOparietal, squamosal, and nasal of S.

terrestris are distinctive. The anterior dorsal portion of

 

the squamosal of S. terrestris is proportionately larger

 

than that of either S. americanus Holbrook or S. woodhousei

 

 

fowleri (Hinckley). The frontoParietal of S. terrestris

 

exhibits a crainial crest which is more rounded and knoblike
than the corresponding member of the latter two species.

The dorsal ridges on the nasal of S. terrestris are more

 

clearly defined than those of S. americanus or S. g.

 

fowleri.

Tihen (1962) records this species from Reddick lB.

Bufo woodhousei fowleri (Hinckley)

 

Material--Eight left ilia, seven right ilia, MSU
643, figures 21, 2L.

Remarks--Fifteen typical EEEQ ilia with ilial
prominence height to base ratios of .30 to .42 are assigned

to Bufo woodhousei fowleri. Although the ilia of Bufo

 

americanus Holbrook also show similar ratios, the base of

 

the ilial protuberance in S. americanus is wider than that

 

of equal sized ilia of S. g. fowleri (Holman, 1967). This
character distinguished the fossils as S. g. fowleri.

Tihen (1962) indicates having seen "one or a few"
ilia from various florida Pleistocene localities that should

be referred to this species. Reddick 13 is mentioned as

15

one of the localities. S. g. fowleri does not occur in

Florida today.

Bufo qpercicus Holbrook

 

Material--One left and one right ilium, MSU 650,
figure 2K.
Remarks--These two small Bufo ilia are difficult to

distinguish from juvenile Bufo woodhousei fowleri

 

(Hinckley). Since they lack the porosity characteristic
of juveniles of the latter, they are assigned to S.

quercicus.

 

Tihen (1962) records this species from the present

site.

Family Hylidae
Hyla cinerea (Schneider)
Material--One left ilium, MSU 647, figure 2N.
Remarks--The large size and oval shape of the dorsal
protuberance, its relatively great distance from the
acetabular fossa, and the relatively acute angle with which
the anterior edge of the ventral acetabular expansion meets

the ilial shaft are characteristic of Hyla cinerea.

 

This is the first fossil record of S. cinerea at
the Reddick 18 site. Lynch (1965) records this Species

from the Pleistocene of Alachua County, Florida.

16

Pesudacris nigrita (Le Conte)
Material--One right ilium, MSU 648, figure 3D.

Remarks--The ilium of Pseudacris nigrita exhibits

 

a wider ventral acetabular expansion than is found in either

S. ornata (Holbrook) or S. triseriata (Wied) and can be

 

separated from those two species on that basis. This
feature is shown in figures BA, BB, and 3C.

This is the first fossil record of this species.

Family Emydidae
Terrapene carolina (Linnaeus)

Material--One proneural, four partial hyoplastrons,
three partial hypoplastrons, two humeri, one scapula, two
femora, MSU 668, figures 3E, 3F.

Remarks--Auffenberg (1958), Holman (1967), and

Milstead (1967), discuss characteristics of Terrapene

 

carolina.
Auffenberg (1959), working with complete shells

from this area, has indicated that the Reddick lB Terrapene

 

probably represents an intergradient population of g. g.
carolina or S. g. bauri and the extinct S. g. putnami
(Hay). However, Blaney (1971) has placed 3. g. putnami

in the snyonomy of g. E. major (Aggasiz).

Gopherus pglyphemus (Daudin)

 

Material--One partial mandible, four humeri, five

femora, MSU 669, figures 36, 3H, 31.

17

Figure 3. Reddick amphibian and reptile fossils.
A. Recent agendacris triseriata ilium, x 6.0.
B. Recent Eggggacris ornata ilium, x 6.0.
C. Recent Psegdacris nigrita ilium, x 6.0.
D. Fossil Pseudacris nigrita ilium, x 6.0.
E. Fossil Terrapene carolina humerus, x 2.0.

F. Fossil Terrapene carolina femur, x 1.5.

 

G. Fossil Gopherus polyphemus humerus, x 2.4.
H. Fossil GOpherus pglyhpemus femur, x 2.0.
I. Fossil Gogherus polyphemus mandible, x 1.4.

J. Fossil Geochelone crassiscutata dermal ossicle, x
3.3.

K. Fossil Anolis carolinensis frontal, x 4.0.

L. Fossil Anolis carblinensis dentary, x 5.0.

 

M. Fossil Cnemidophorus sexlineatus dentary, x 5.0.

 

N. Fossil Ophisaurus ventralis precadal vertebra, x
4.0.

 

0. Fossil Ophisaurus ventralis caudal vertebra, x 3.8.

P. Fossil Ophisaurus ventralis dentary, x 5.0.

 

Q. Fossil Eumeces, Sp. indet. dentary, x 5.0.
R. Fossil Elaphe, sp. indet. verbetra, lateral, x 4.0.
S. Fossil Elaphe, sp. indet. vertebra, dorsal, x 4.0.

T. Fossil Elaphe, sp. indet. vertebra, ventral, x 4.0.

18

 

 

Q

Figure 3. Reddick amphibian and reptile fossils.

19

Auffenberg (1959) records Gopherus from this site.

Order Testudines
Family Testudinidae

Geochelone, cf. S. crassiscutata (Leidy)

 

Material--One dermal ossicle, MSU 670, figure 3J.
Remarks--The size of the ossicle (3 cm.) indicates

the presence of a large tortoise. Only S. crassiscutata

 

is known to reach such a size in the Pleistocene of
Florida.

Auffenberg (1963b) records this species from this

site.

Geochelone, cf. S. incisa (Hay)
Materia1--Fragmented carapacial elements, MSU 671.
Remarks--Several carapacial elements, too
fragmentary for positive specific identification
exhibit the rugosity and thickness characteristics of
this species.
Auffenberg (1963b) records this species from this

site.

Order Squamata
Family Iguanidae

Anolis carolinensis Voigt

 

Materia1--Five dentary fragments, five frontals,

MSU 652, figures 3K, 3L.

20

Remarks-—Anolis dentaries are easy to identify.

 

They are long, low, and have a reduced or absent Meckelian
Groove. The heterodont teeth are singly cusped anteriorly,
tricusped posteriorly. I cannot distinguish the fossils
from recent Anolis carolinensis dentaries.

The frontal of Anolis is also distinctive. The

fossil frontals are assigned to S. carolinensis.

 

Auffenberg (1956) records S. carolinensis from

 

Reddick 18.

Family Teidae
Cnemidophorus sexineatus (Linnaeus)

Material--Three dentaries, two vertebrae, MSU 653,
figure 3M.

Remarks--The fossils are indistinguishable from
corresponding elements of Cnemidophorus sexlineatus. On
this basis and on zoogeographic grounds, the fossils are
assigned to that species.

Auffenberg (1956) records a single dentary fragment

from Reddick 13.

Family Anguidae
Ophisaurus ventralis (Linnaeus)
Material--Eight precaudal vertebrae, five caudal
vertebrae, MSU 655, figures 3N, 30, 3P.
Remarks--Auffenberg (1955) and Etheridge (1961)
have discussed methods for identifying the vertebrae of the

three species of Ophisaurus occurring in eastern North

 

21

America. They have found that precaudal vertebrae of S.

compressus Cope can be distingushed from those of S.

 

attenuatus Baird and S. ventralis by the shorter and wider

 

 

appearance of the latter two Species. S. attenuatus

 

precaudal vertebrae can be separated from those of S.

ventralis by using the angle formed by the posterior border

 

of the neural spine and the longitudinal axis of the
centrum. These angles have been reported to be 45° to 65°

(mean--58°) for S. attenuatus, and 65° to 84° (mean-~73°)

 

for S. ventralis. The angle between the anterior border

 

of the caudal transverse processes and the longitudinal
axis of the centrum is used to diagnose the caudal
vertebrae of all three Species. These angles have been

reported to be 70° to 75° (mean--73°) in S. ventralis,

 

75° to 85° (mean--81°) in S. attenuatus, and 50° to 65°

 

(mean--55°) in S. compressus.

 

Using the criteria given above, the fossils were

identified as S. ventralis. In the course of identifying

 

the fossils, four prepared skeletons of S. attenuatus

 

attenuatus were examined. The neural Spine-longitudinal

 

axis angle in these Specimens varied from 68° to 81°
(mean--72°) thus causing the known variation of this
measurement to overlap with the known variation in S.

ventralis. Assuming that Etheridge and Auffenberg used the

 

eastern subSpecies of S. attenuatus (S. 3. longicauda) as

 

 

their reference material, it may be possible to use

vertebral angles to separate S. ventralis from

 

22

S. attenuatus longicauda. Making this assumption these

 

fossils are identified as S. ventralis.

 

Auffenberg (1955) records S. ventralis from this

 

site.

Family Scincidae

Eumeces, sp. indet.

Material--Two partial dentaries, MSU 654, figure 3Q.

Remarks--Auffenberg (1956) provisionally lists
Eumeces fasciatus (Linnaeus) from Reddick 18. His identi-
fication was based on a single complete dentary containing
twenty-seven teeth. The recent skeletons available to him
gave dentary teeth counts of twenty-one to twenty-five in

Eumeces laticeps Schneider and Eumeces inegpectatus Taylor,

 

and twenty-five to twenty-six in S. fasciatus. Since.the

 

number of teeth found in S. fasciatus most closely

 

approached the number found in the fossil, the fossil was
assigned to that Species.

Two additional fossil dentaries from the Reddick
13 site are assigned to Eumeces. One fossil dentary is
nearly complete and has a tooth-alveolar count of twenty-

four, placing it in the laticeps and inexpectatus group of

 

Auffenberg. However, eight MSU skeletons of S. fasciatus

 

yielded tooth counts of twenty—three to twenty—four,
indicating that the fossil might represent any of those

three species.

23

The other fossil dentary is too fragmentary for a
complete tooth count. I am unable to carry this identifi-

cation of either fossil dentary to the Species level.

Family Colubridae

Carphophis amoenus (Say)

 

Material--Eleven precaudal vertebrae, MSU 660.

Remarks-~Diad0phis, Tantilla, and Rhadinea are

 

other small colubrids whose vertebrae might be confused with
those of Carphophis. Auffenberg (1963) presented data

showing the neural Spine of Carphohpis vertebrae is

 

proportionately longer and lower than that of Diadophis or

 

Rhadinea. He further indicates that Tantilla vertebrae

have a narrower haemal keel than those of CarphOphis.

 

Using these criteria, the fossils were identified as

Carphgphis amoenus.

 

Elaphe, Sp. indet.

Materia1--Twenty-seven precaudal vertebrae, MSU
659, figures 3R, 38, 3T.

Remarks--Auffenberg (1963b) found the reference

skeletons of Elgphe guttata (Linnaeus) and Elaphe obsoleta

 

 

(Say) available to him could be separated by a regression
curve of ratios of the neural Spine length to neural spine
height. Enough overlap in this character has been found in
MSU Specimens to invalidate its usefulness in diagnoses.

I am unable to separate these two species by vertebra

characters.

24

Pituophis melanoleucus (Daudin)

 

Material--One Sphenoid, twenty precaudal vertebrae,
MSU 657, figures 4C, 4D, 4E.

Remarks--The broad, short appearance of the
sphenoid makes it distinctive. The proportionately high

neural spine of eastern S. melanoleucus serves to distinguish

 

it from other large colubrids. From Elaphe with which it is
most easily confused it can be distinguished its prOpor-
tionately larger neural canal and the more vaulted
appearance of its neural arch. Figures 4A and.4B illustrate

these differences.

Coluber or Masticophis

 

Material-~Thirty—one precaudal vertebrae, MSU 658,
figures 4F, 4G.
Remarks--I am unable to distinguish between the

vertebrae of Coluber constrictor Linnaeus and MasticoPhis

 

 

flagellum (Shaw), the two extant members of these genera

 

in Florida.

Drymarchon corais (Daudin)

 

Material--One maxilla, one palatine, one basis-
phenoid, one articular—surangular-prearticular complex,
thirty—one precaudal vertebrae, MSU 656, figures 4H, 41,
4J, 4K.

Remarks--The crainial elements are distinctive and

not easily confused with other Species. The vertebrae are

Figure 4.

A.

B.

C.

J.

K.

L.

Recent

Recent
x 3.2.

Fossil

Fossil
2.5.

Fossil
x 2.5.

Fossil
x 2.6.

Fossil
x 3.0.

Fossil

25

Reddick snake fossils.

Elaphe obsoleta verbetra, posterior, x 3.2.

 

Pituophis melanoleucus vertebra, posterior,

 

Pitu0phis melanoleucus sphenoid, x 3.0.

 

Pituophis melanoleucus verbetra, dorsal, x

 

Pitu0phis melanoleucus vertebrae, lateral,

 

Coluber or Masticgphis vertebra, lateral,

 

Coluber or Masticophis vertebra, dorsal,

 

Drymarchon corais maxilla (upper), palatine

 

(lower), x 2.7.

Fossil
Fossil
Fossil
Fossil

Fossil
x 4.0.

Fossil

Drymarchon corais sphenoid, x 1.9.

 

Drymarchon corais vertebra, lateral, x 2.0.

 

Drymarchon corais vertebra, dorsal, x 2.1.

 

Heterodon platyrhinos Sphenoid, x 4.0.

 

Heterodon platyrhinos vertebra, anterior,

 

Heterodon Simus vertebra, anterior, x 4.0.

 

26

 

Reddick snake fossils.

Figure 4.

27

easily identified by the beveled anterior edge of their
neural spine (Auffenberg, 1963b).

All of these bones were found together in one
shovelful of matrix. The size of the various elements

indicates that they all may have come from one adult snake.

Heterodon platyrhinos Latreille

 

Material--One sphenoid, twenty-one verbebrae, MSU
661, figures 4L, 4M.

Remarks--The sphenoid is dinstinctive. Its
pleurosphenoid portion is more convex in outline than any

other colubrid of the southeastern United States. Heterodon

 

vertebrae are easily identified by their flattened and

broadened haemal keel (Holman, 1959).

Heterodon simus (Linnaeus)

 

Materia1--One precaudal vertebra, MSU 662, figure
4N.
Remarks.--In addition to the characters used by

Auffenberg (1963b) to separate S. simus from S. platyrhinos,

 

the shape of the neural canal can be used also. In S.
simus the canal is more dorsally expanded than in S.

platyrhinos.

 

Storeria dekayi (Holbrook)

 

Materia1—-Two precaudal vertebrae, MSU 663.

28

Remarks--The two fossils appear to be middle
precaudal vertebrae. Using the criteria of Auffenberg

(1963b), the fossils are assigned to S. dekayi.

Thamnophis sirtalis (Linnaeus)

 

Material--One sphenoid, sixty-three precaudal
vertebrae, MSU 664, figures 5E, 5F, 5G.

Remarks--The sphenoids of various large natricines
are very similar. Holman (1967) points out that Sphenoids
of S. sirtalis have more pointed, better develoPed posterior

processes than those of Natrix sipedon (Linnaeus) and S.

 

Sguritus (Linnaeus). The fossil Sphenoid is similar to S.
sirtalis in this character and are assigned to that species.
Holman (1962b) and Auffenberg (1963b) have dis-

cussed the vertebral characteristics of Thamnophis. As

 

they point out, the large natricines with which Thamnophis

 

vertebrae are most easily confused are Natrix sipedon

 

(Linnaeus) and S. fasciata (Linnaeus) both of which have
higher neural spines than S. sirtalis. Figures 5A, SB, and
SC illustrate this point.

Besides the characters mentioned by Holman (1962b),
S. sirtalis vertebrae can be separated from those of S.
sauritus by the lower neural spine and prOportionately

wider neural arch of the latter.

Figure 5.

A.

B.

C.

D.

E.

F.

G.

H.

Recent
Recent
Recent
Recent
Fossil
Fossil
Fossil
Fossil
Fossil
Fossil
Fossil

Fossil
x 3.7.

Fossil
Fossil

Fossil
x 2.0.

29

Reddick snake fossils.

Natrix fasciata vertebra, lateral, x 4.0.

 

Natrix sipedon vertebra, lateral, x 4.0.

 

ThamnOphis sirtalis vertebra, lateral, x 4.0.

 

Thamnophis sauritus vertebra, lateral, x 4.0.

 

ThamnoPhis sirtalis sphenoid, x 4.0.

 

Thamnophis sirtalis vertebra, lateral, x 3.6.

 

ThamnoPhis sirtalis vertebra, dorsal, x 3.0.

 

Micrurus fulvius vertebra, lateral, x 3.6.

 

Micrurus fulvius vertebra, dorsal, x 3.9.

Sistrurus miliarius vertebra, lateral, x 3.1.

 

Sistrurus miliarius vertebra, dorsal, x 4.9.

 

Crotalus or Sistrurus fang and premaxilla,

 

Crotalus adamanteus vertebra, lateral, x 2.0.

 

Crotalus adamanteus vertebra, dorsal, x 1.8.

 

Crotalus adamanteus vertebra, anterior,

 

 

30

 

Reddick snake fossils.

Figure 5.

31

Family Elapidae

Micrurus fulvius (Linnaeus)

 

Material-~One-hundred sixty-six precaudal vertebrae,
MSU 665, figures 5H, SI.

Remarks-~These are typical Micrurus vertebrae with
well developed hypapOphyses, and a long, low neural spine.
0n zoogeographic grounds they are assigned to S. fulvius
from which they cannot be distinguished.

This was the single most common reptilian element
found in the samples. Whether the vertebrae represent few

or many individuals is uncertain.

Family Viperidae

Sistrurus miliarius (Linnaeus)

 

Material--Thirty-five precaudal vertebrae, MSU 666,
figures SJ, 5K.

Remarks--The fossils are typical Sistrurus

 

vertebrae, being of the viperid type and having a relatively
long and narrow centrum. On zoogeographic grounds they are

assigned to S. miliarius rather than S. catenatus.

 

 

One viperid maxilla with its attached fang was
found. Its small size suggests that it may represent this
Species. But, I am unable to distinguish between S.
milarius and Crotalus adamanteus Beauvois using this ele-
ment. The squarish outline of the maxilla indicates that

the fossil does not represent Agkistrodon. Holman (1959),

 

32

has pointed out that Sgkistrodon maxillae have a more

rounded outline. This fossil is illustrated in figure 5L.

Crotalus adamanteus Beauvois
Material-~Ten precaudal vertebrae, MSU 667,
figures 5M, SN, 50.
Remarks-~Auffenberg (1963b), discusses the vertebral

characteristics of S. adamanteus. Several of the vertebrae

 

represent large individuals, but they do not exhibit the

wide prOportions of the extinct S. giganteus Brattstrom.
DISCUSSION

The Herpetofauna
In their check list of 1963, Gut and Ray list
thirty-eight species of reptiles and amphibians as being
found at Reddick I. With the exception of Pseudobranchus
robustus Goin and Auffenberg, and Rhineura floridana
Baird and Girard, all of the species listed by Gut and Ray
were found at the 13 site. Holman (1962a) originally

recorded S. robustus and S. floridana from Reddick IC.

 

The present study adds five species to the known
herpetofauna of Reddick IB, Ambystoma tigrinum, Rana ggylio,

Rana areolata, Hyla cinerea, and Pseudacris nigrita. This

 

is also the first fossil record of Rana areolata and

 

Pseudacris nigrita.

 

The herpetofauna from the Reddick I site is com-

prised of at least forty-three Species, of which all but

33

two occur at the IB site. S. robustus, S. crassiscutata,

 

and S. incisa are the only extinct forms. Thus approxi-
mately 40 percent of the ninety-three extant Species of
reptiles and amphibians found in central Florida are

represented in the Reddick fossil fauna.

Paleoecology

Although it is generally agreed that the Reddick l
fossils accumulated in a large cave (Brodkorb, 1957;
Auffenberg, 1963b, Hamon, 1964), various investigators have
expressed divergent Opinions as to the nature of the
surrounding paleocommunity. Based on the avifauna, Brod-
korb (1957) and Hamon (1964) suggest that a typical wet
grassland or freshwater marsh was nearby. Brodkorb (1957)
lists the following species of fossil birds from Reddick l
as being characteristic of such a habitat: piedbilled grebe,
six species of duck, six species of rails, killdeer, lesser
yellowlegs, burrowing owl, sedge wren, yellow throat, red-
winged blackbird, meadowlark, and Henslow's Sparrow. He
also notes that the remaining avian members of the fossil
fauna are in harmony with a marsh community. The absence
of birds which are confined to deep water and the total
absence of fish remains were interpreted as meaning that the
marsh contained no sizeable body of open water and that the
marsh probably became dry periodically.

Auffenberg (1963) interprets the reptilian faunal

elements as suggesting a dry, open forest, but not

34

necessarily scrub. He points out that the absence of
aquatic genera such as Natrix suggests that no ponds were
nearby. He further notes that the bulk of Brodkorb's bird
fossils came from site 1A, while the reptiles (primarily
snakes) came from site 13 which is situated approximately
260 feet to the southwest. This leads him to speculate that
these two sites may not be contemporaneous, and may”
represent different ecological conditions.

The fossil herpetofauna being discussed in this
paper was taken from one small area of locality 13. The
possibility that this sample represents a heterochronic
assemblage rather than a unit fauna still exists, but that
likelihood is reduced due to the small volume of the sample
and the localized nature of the sampling technique.

Most of the Species found in the present sample
have wide ecological tolerances and may be found in a
variety of habitats. It is the species whose habitats are
restricted that give the best clues to the paleohabitat.

In the latter category are Siren lacertina, Pituophis

 

 

melanoleucus, and Gopherus pglyphemus. S. lacertina

 

requires an aquatic habitat, but can estivate in moist
burrows during droughts (Carr, 1940). S. melanoleucus and

S. polyphemus are largely restricted to high pine, although

 

S. polyphemus is occasionally found in upland hammock

 

situations (Carr, 1940). Thus in this one sample elements

are present which are strong indicators of the dry Open

35

forest postulated by Auffenberg and the freshwater marsh
of Brodkorb.

Of the remaining members of the fauna, Carr (1940)
characterizes twelve as being characteristic of or occurring
frequently in high pine, while seven are listed as being
characteristic of or occurring frequently in upland
hammocks (see Table 1). Nearly all of the fauna can be
found at least occasionally in either the high pine or
upland hammock habitats. The addition of a nearby marsh to
either habitat would allow for the presence of the aquatic
members of the fauna.

It is entirely possible that a high, well-drained
forest could be associated with a lower marsh area. Such
conditions are found in central Florida today. Hamon
(1964) states that the mean surface elevation of Reddick l
is ninety-three feet. He also furnishes a relief map of
this area which indicates the presence of a shallow
basin. The center of this basin lies within one thousand
feet of the Reddick l quarry and is at an elevation of
seventy feet. Some of the terraine surrounding the basin
and quarry rises to an elevation of 120 feet. Under
suitable conditions the basin could become a shallow pond
with a radius of several hundred feet.

Considered as a unit fauna, the sample suggests
that both Auffenberg and Brodkorb's views of the paleo-
ecology of the Reddick 1 site may be partially correct. A

synthesis of their two paleoecological interpretations seems

36

Table l.--Habitat preferences of the Reddick lB herpeto-
fauna.

 

Species High Pine Upland Hammock Marsh

 

Siren lacertina - - c

 

Ambystoma tigrinum -

 

O O
O

Scaphiopus holbrooki f
Rana grylio
Rana pipiens

Rana areolata

 

I
I
O

 

 

Gastrophyrne carolinensis
Bufo terrestris

 

Bufo S. fowleri
Bufo quercicus

 

Syla cinerea

 

Psuedacris nigrita

 

Terrapene carolina

 

Gopherus polyphemus

 

Anolis carolinensis

 

Cnemidophorus sexlineatus

HHOOOOOHOHOO

Ophisaurus ventralis

Carphophis amoenus
Pituophis melanoleucus

 

Coluber or Masticophis

 

Drymarchon corais

 

Heterodon platyrhinos

 

OHIOH’tO

Heterodon simus

 

Storeria dekayi

 

Thamnophis sirtalis

 

Micrurus fulvius

 

Sistrurus miliarius

 

OOHOOOOOOOOOOHOFHOOOOOHIOO

HiOOO

Crotalus adamanteus

 

 

Note: f = frequent; c = characteristic; 0 =
occasional; - = rare or absent. Data from Carr (1940), and
Conant (1958).

37

to be the most satisfactory explanation of the apparent
ecological incompatibility of the species found at

Reddick l.

Zoogeography and Pleistocene Climate

Were it not for the presence of S. crassiscutata,

 

S. S. fowleri, and S. amoenus, the fossil herpetofauna of

Reddick 18 would offer no suggestion that the environmental
conditions under which they lived differed in the least

from the present conditions in that area. However, S. S.
fowleri and S. amoenus presently occur no farther south than
several hundred miles to the north of Reddick (figure 5).

S. crassiscutata, a giant tortoise, is now extinct. The

 

giant tortoise which occurs closest to the Reddick site is
found in the Galapagos Islands. This information implies
that the ecological relationships between the faunal
members, and possibly the existing climatic conditions,
differed from current conditions.

Several authors have given their interpretation of
the paleoclimate at Reddick l. Brodkorb (1957), believing
that the Reddick 1 fauna represented the Illinoian glacial
stage, suggested that when the fossils were deposited the
climate of northern Florida was at least as cool as that
of northern Virginia. His conclusion was based on finding

the bones of nestling Tachcineta speleogytes Brodkorb, an

 

extinct swallow. The closest living relative of S.

Speleodytes breeds no farther south than Virginia. Among

38

 

 

 

I-d

/'\-+F\ 1”

 

 

 

 

 

   

s... 8%

 

 

 

Figure 6. Southern range limits of CarphOths amoenus
and Bufo EI- fowleri.

---- S. amoenus

..... B. S. fowleri

39

the fossil mammals found at Reddick l, Brodkorb points to

Synaptomys australis and notes that today the genus

 

Synaptomys finds its southern limits in Maryland, although

 

isolated populations occur in Virginia and North Carolina.
Hamon (1964) attempted to determine the July and

January mean temperatures at Reddick at the time of fossil

preservation. He mapped the present southern limits of

Synaptomys, Tachycineta, and Cistothorus, three boreal

 

 

elements found in the deposit of Reddick. An average July
mean temperature of 74.9 degrees Fahrenheit was found to
occur along these borders. That temperature was compared
with a July mean temperature of 81.1 degrees Fahrenheit at
Ocala, Florida, a location ten miles south of the fossil
site. Hamon concluded that during the time of fossil
deposition the summers at Reddick were about six degrees
cooler than now.

To determine the January mean temperature the
process was repeated using the northern range of a southern

element. Alligator mississipiensis was chosen as this

 

southern element. The average January mean temperature
along the northern limits of its range was found to be 50.4
degrees Fahrenheit. This temperature was equated with the
January mean temperature of Reddick at the time the fossils
were being deposited. The current January mean temperature
at Ocala, Florida was stated to be 57.2 degrees Fahrenheit,
about seven degrees warmer than Reddick at fossil deposition

time. Thus, both Brodkorb and Hamon concluded that at the

40

time the fossils were being deposited, summer and winter
temperatures were lower than now. These conclusions and
the methods used to arrive at them are open to criticism.

Loosely paraphrased, Merriam's temperature laws
state that animals and plants are restricted in northward
distribution by the total quantity of heat during the season
of growth and reproduction, and that their southward
distribution is determined by the intensity of heat during
the hottest period of summer. Either knowingly or un-
knowingly, both Brodkorb and Hamon used these laws, or a
facsimile thereof, to arrive at an estimation of the
paleoclimate at Reddick. Since these laws have been largely
discredited (Kendeigh, 1932; Shelford, 1932), their use in
the determination of paleoclimates must be questioned.

The use of the alligator's northern range limits to

estimate paleotemperatures at Reddick must be questioned,
also. The present range of the alligator extends to the
north of Reddick (Conant, 1958, Map 1). Therefore, it is
meaningless to equate the average the January mean temper-
atures along its northern limits with the January mean
temperature of Reddick at the time of fossil deposition.
To arrive at an estimation of the January mean at Reddick
in a manner analogous to that which was used to estimate
the July mean temperature, a southern element should have
been chosen whose northern range now lies to the south of
the Reddick site. This might be possible to do if the

January paleotemperatures were warmer than at present. But

41

again, the practice could be criticized due to the
questionable validity of Merriam's temperature laws.

In recent years, other workers (Dalquest, 1965;
Hibbard, 1970; Slaughter & Hoover, 1963) have made inter-
pretations of Pleistocene paleoclimates of the United
States. Their works have centered on the Great Plains

region, and have been based largely on faunal evidence.

In particular, fossils representing species having northern
distributions have been used as temperature indicators.
When such a fossil was found to the south of the current
southern range limits of the species it represented, it was
interpreted as an indication that cooler summer temperatures
prevailed at the paleohabitat. As already discussed, such
interpretations are loosely based on Merriam's temperature
laws, and therefore, they are based on faulty assumptions.
Nevertheless, the importance of temperature effects
cannot be overlooked as a possible factor, perhaps even the
major factor, in contributing to faunal mixing as found in
Pleistocene deposits. However, other factors may also
contribute to the phenomenon. Humidity, seasonal patterns
of rainfall, predation, and competition are just a few of
the factors that may act independently or in concert with
other environmental factors to delimit a species' range.
As yet, just how these factors may interact to affect
distributions is only partly understood. Until such inter-

actions are more fully investigated, it seems unwarranted

42

to ascribe all faunal mixing to temperature effects alone,
as has been done.

Therefore, although the presence of g, g. fowleri
and g. amoenus does not conflict with the hypothesis that
cooler summer temperatures prevailed at Reddick during the
time of fossil deposition, neither does it offer positive
evidence for it. Clark (1970), studying the natural
history of Carphophis in detail, attributes the occurrence
of Carphophis in Pleistocene Florida to the better soil
drainage that was presumed to be present then. The
same or a similar non—temperature factor may account
for the presence of E. w. fowleri, also.

The foregoing discussion might seem to make an
estimation of paleotemperatures based on faunal elements
impossible. This is not necessarily the case. If it can
be shown that a particular species' distribution is deter-
mined entirely or largely by temperature, then that species
could be used as a temperature indicator. Some progress
has been made in this direction.

Brattstrom (1961) notes that mild, equable climates
are necessary for the existence of giant tortoises. With-
out such climatic conditions and some shade, the tortoises
reach lethal temperatures when exposed to the sun. In a
cool climate their great bulk takes too long to warm up,
and they cannot become active. In addition, Brattstrom
shows that tortoises have always have a tropical or

subtropical distribution.

43

Hibbard (1960) also suggests that giant tortoises
are good indicators of equable climates. He notes that
specimens of the giant tortoises brought from the Galapagos
Islands were not able to survive in colonies near the
southern border of the United States. Even specimens
located on Lignum Vitae Island, Florida, died from enteritis,
a condition caused by the tortoises being too cool to
digest their food.

Hibbard's use of the giant tortoises as indicators
of climate is conservative. His summarizing statements on
their use for this purpose bears repeating.

The large Pliocene and Pleistocene land tortoises
could not stand freezing, but they may have existed in
an area where very few light frosts occurred at night
but with temperature during the day warmed to 60
degrees or more Fahrenheit.

The young of these large species may have taken
shelter on cool nights in burrows until they were
large enough so that their body heat would protect
them from temperatures near 32 degrees F. The fact
that there is no evidence that these tortoises burrowed
or entered burrows during winter months, and the fact
that they grew to such large size, suggests that they
lived and reproduced in an equable climate.

It is therefore assumed that they lived chiefly
in areas of frost free environments. It is not
logical to assume that these large land tortoises
lived in an environment where they became torpid from
cold and dotted the landscape as boulders through a
long cold season (Hibbard, 1960, p. 9).

Geochelone crassiscutata is a giant tortoise whose
remains have been found at the Reddick 18 site. Auffenberg
(1963b) records a specimen of this species whose carapace
measured 1210mm in total length. He also states that

fragments of much larger specimens have been found.

44

The presence of g. crassiscutata in the Reddick

 

fauna suggests that when this species lived there, the
climate was equable and frost free. This would seem to
indicate that winter temperatures were not quite so cold as

they are now. Data taken from the United States Weather

 

Bureau Climatic Summary Supplement, 1932-62, showed that

 

at Ocala, Florida, the temperature dr0ps below thirty-two
degrees Fahrenheit on an average of twelve days per year.
Presumably, this would make the area uninhabitable by the

giant tortoise.

SUMMARY

Five species have been added to the faunal list of
fossil reptiles and amphibians found at the Reddick 1 site.
The total recorded herpetofauna now stands at forty-four
species, including three extinct species.

The fossils identified at site 13 indicate that
the surrounding area once contained a well-drained, open
forest and a marsh.

The presence of the extinct giant tortoise

Geochelone crassiscutata suggests that the winters at

 

Reddick during the Sangamon Age were not as cold as

present.

PART II
THE PLEISTOCENE HERPETOFAUNA OF

LADDS, GEORGIA

The only site between north-central Florida and
north-central Tennessee which has yielded an extensive
Pleistocene herpetofauna is found near Ladds, Georgia.

The Ladds deposit is in a limestone quarry located 2.3
miles west south-west of Cartersville, Bartow County, in
Land Lot 591, Fourth District, Third Section of the
Georgia Land Survey of 1832. The quarry is developed at
the southeastern end of an isolated dolomite ridge, Quarry
Mountain. The ridge stands about five hundred feet above
the surrounding countryside and reaches an elevation of
nearly eleven hundred feet.

Solution cavities of the dolomite contain flowstone
and red cave earths which may surround or encrust the
fossils. Due to blasting and other quarrying operations,
ample opportunity for the mixing of heterochronic elements
has occurred (Ray, 1965; Lipps & Ray, 1967).

Faunal evidence suggests the deposit is late
Pleistocene. Ray (1967), reported that nearly 25 percent

of the mammals found in the deposit are extinct. He

45

46

reports the presence of late Pleistocene indicators such

as Equus and Didelphis.

 

For the most part, collecting of fossils has been
done by personnel affiliated with Shorter College, Rome,
Georgia, under the direction of Dr. Lewis Lipps. Standard
collecting techniques have been used, including washing for
microfossils.

Beginning in 1963, fossils were collected, sorted,
and shipped from Shorter College to the Smithsonian Insti-
tution. There, Dr. Clayton Ray separated the material
into various taxonomic classes and shipped the reptile and
amphibian portion to Dr. J. Alan Holman for study. Holman
(1967) reported on the fossils which were collected in
1963 and 1964 and were studied by him.

Since then, a wealth of new reptile and amphibian
fossils from the Ladds site have been received for study.
These fossils, dated pre-l968, 1968, and 1969, are the
subject of this account.

The following is a list of the reptiles and
amphibians represented by the fossils identified in the

samples labeled pre-l968, 1968, and 1969.

Class Amphibia
Order Urodela

Family Ambystomatidae
Ambystoma tigrinum (Green)
Family Plethodontidae
Plethodon cf. Plethodon glutinosus (Green)

 

47

Order Anura

Family Ranidae

Rana i iens Schreber
Family Bufonidae

Bufo americanus Holbrook
Bufo terrestris (Bonnaterre)

 

 

Class Reptilia

 

Order Testudines

Family Emydidae
glemmys insculpta (Le Conte)
Terrapene carolina (Linnaeus)
Chrysemys picta—TSchneider)
Pseudemys sp. indet.

 

 

 

 

Order Squamata

Family Colubridae
QiadOphis punctatus (Linnaeus)
Carphophis amoenus (Say)
Coluber or Masticgphis
EIaphe sp. indét.
Lampropeltis getulus (Linnaeus)
LamprOpeltis triangulum (Lacepede)
gatrix sipedon (Linnaeus)
Storeria dekayi (Holbrook)
Thamnophis sirtalis (Linnaeus)
Family Viperidae
Crotalus horridus (Linnaeus)

 

 

 

ANNOTATED LIST

Class Amphibia

 

Order Urodela
Family Ambystomatidae

Ambystoma tigrinum (Green)

 

Material--Eight vertebrae, USNM 209175.

Remarks--These Ambystoma vertebrae are assigned to

 

A. tigrinum since the posterior portion of the neural arch

of each is upswept and extends well beyond the end of the

48

centrum. Holman (1969) has found this character to be
peculiar to A. tigrinum.

As pointed out by Holman (1967) A. tigrinum does
not occur in north-western Georgia today, although it is

found in nearby southeastern Tennessee.

Family Plethodontidae

Plethodon, cf. Plethodon glutinosus (Green)

 

 

Material--Fifty—two vertebrae, USNM 209176.

Remarks--P1ethodon glutinosus vertebrae are

 

 

amphicoelus. They can be separated from the vertebrae of
Leurognathus and Desmognathus which are Opisthocoelous, and

from Pseudotriton and GyrinOphilus vertebrae which are

 

falsely Opisthocoelous (Soler, 1950). Holman (1969)
separates Plethodon glutinosus vertebrae from those of

Eurycea bislineata (Green), E. longicauda (Green), and g.

 

 

lucifuga (Rafinesque) on the basis of the shorter and wider
dimensions of the former. He also notes that adult g.

glutinosus vertebrae are larger than those of Plethodon

 

 

cinereus (Green), E. bislineata and the troglodyte species

 

of Eurycea. Using these criteria, the fossil vertebrae are

assigned to Plethodon glutinosus from which they cannot be

 

separated.

 

49

Order Anura
Family Ranidae
3233 pipiens Schreber .

Haterial--Twelve right and six left ilia, USNM
209177.

Remarks--Auffenberg (1957) separates §§g§_ilia into
two groups. The pipiens group is characterized by a less
acute angle formed by the posterodorsal border of the idial
crest and the dorsal acetabular expansion. The fossils
represent members of the pipiens group. Holman (1967)

separated g. pipiens from 3. palustris Le Conte on the

 

basis of the former's steeper slope of the posterodorsal
border of the ilial shaft into the dorsal acetabular
expansion. He further separated 5. pipiens from 3.
sylvatica Le Conte in that the former has a larger, less
produced and less roughened prominence for the origin of
the vastus externus head of the triceps femoris muscle.

The fossil ilia agree with 5. pipiens in these characters.

Family Bufonidae

Bufo americanus Holbrook

 

Materia1--Three left and three right ilia, USNM
209178, figure 7a.
Remarks—~All six fossil ilia are much larger than

ilia of Bufo quercicus Holbrook, and they lack the charac-

 

teristic ilial protuberance of that species. However, the

50

Figure 7. Ladds amphibian and reptile fossils.
A. Fossil Bufo americanus ilium, x 3.7.
B. Fossil Terrapene carolina carapace, dorsal, x 0.6.

C. Fossil Terrapene carolina carapace, lateral x 0.6.

 

Figure 7. Ladds amphibian and reptile fossils.

52

fossil ilia have a better developed ilia protuberance than
is found in 2252 terrestris (Bonnaterre).

The fossils are difficult to separate from.§g£g.
woodhousei fowleri (Hinckley) and §g£g_americanus.
However, as Holman (1967) points out, the base of the ilial

protuberance is wider in B. americanus than in equal sized

 

ilia of B. E. fowleri. The fossils agree with B. americanus
in this character.

Holman (1967) comments on the large size of the
fossils of this species from this same site. He points out
that one of the fossils represent a specimen that exceeds
the size of the largest recorded B. americanus. Two of the
ilia examined in the present study of the Ladd's site match
or slightly exceed the size of the large ilium examined by
Holman. However, except for their large size, these ilia
do not appear to differ from g. americanus in any respect,

and therefore cannot be assigned to a new species.

Bugg terrestris (Bonnaterre)

Material--Five right and four left ilia, USHM
209179.

Remarks--The.ilia of £252 terrestris are easily
separated from Bungamerican, B. quercicus Holbrook, and

g. g. fowleri. g. terrestris has a low ilial prominence

 

with a wide base. The other three species of Bufo have

higher, narrower based ilial prominences.

53

This is the first record of this species from the

Ladds site.

Class Reptilia
Order Testudines
Family Emydidae
Clemmys insculpta (Le Conte)
Material--One right epiplastron, USNM 209180.

Remarks--The epiplastron of Clemmys insculpta is

 

distinct from other North American emydid turtles. Besides
being characteristically narrow and thick, it has a
characteristic deep gutter next to the gular lamina

(Holman, 1967).

Terrapene carolina (Linnaeus)

Haterial--Two right and one left femora, a fusion
consisting of a partial proneural, several partial
peripherals, and one.partial pleural, USHH 209181; one
nearly complete carapace, USNH 26095, figures 7b, 7c.

Remarks--Holman (1967) separates Terrapene femora
from other North American.emydid turtles by their stouter,
more bowed appearance. The fossil femora are similar in

those characters to living Terrapene carolina from which I

 

am unable to separate them. The fossil consisting of the
fused anterior carapacial elements cannot be separated from
Terrapene carolina, either. Therefore it is referred to

that species.

54

Of considerable interest is the nearly complete
carapace of T. carolina. Only a small portion of the pro-
neural bone is missing from this fossil. Today, northern

Georgia is inhabited by T. carolina carolina, but the Ladds

 

site is only several hundred miles removed from the range
of T. g. bauri Taylor, T. 9. major (Agassiz) and T. g.
triunguis (Agassiz). Of these subspecies, the fossil most

closely resembles T. g. carolina and T. g. triunguis. The

 

fossil lacks the characteristic hump behind the third

vertebral scute of triunguis and it does not show the high

 

degree of vaulting often found in triunguis. However, both

 

these characters may be lacking in triunguis and thus the

 

fossil cannot be referred to T. g. carolina with assurance.

I prefer to refer to it as Terrapene carolina, subspecies

 

indeterminate.

Chrysemys picta (Schneider)
Material--One left epiplastron, USNM 209182.

Remarks--The epiplastron of Chrysemys picta is

 

easily distinguished from other North American emydid
turtles. It differs from Clemmys species in being broader
and thinner. It is broader than Deirochelys reticularia.

It differs from Emydoidea in having a well developed,
laterally directed process immediately anterior to the gular
lamina. In T. 22533 the gular and abdominal scutes cover a
large portion of the dorsal surface of the epiplastron, and

produce a wide flat border on it. This wide, flat border

55

is lacking in Malaclemmys, Graptemys species, and Pseudemys
species. The fossil epiplastron possesses this character

and is hence referred to 9. pi t .

Pseudemys, sp. indet.

 

Material-~One right partial hypoplastron USNM
209183.

Remarks--This large fossil hypOplastron has the
general conformations of a large Pseudemys, but because of
its fragmentary nature I am unable to carry the identifi-
cation to the species level.

Holman (1967) described a large fossil proneural
from this site. He provisionally referred the specimen to

Pseudemys concinna (Le Conte) and made note of its large

 

size. He estimated that the fossil proneural represented

a turtle with a carapace length of at least fourteen inches.
The fossil hypoplastron is five inches long

(127 .0 mm). A turtle of the Pseudemys group would have

a corresponding carapace length of at least sixteen inches.

Conant (1958) records only g. concinna, g. floridana, and

 

g. rubriventris as approaching or exceeding this size.

 

Order Squamata
Family Colubridae

DiadOphis punctatus (Linnaeus)

 

Material--Twelve precaudal vertebrae, USNM 209184.
Remarks--Holman (1967) remarks on characters which

separate DiadOphis punctatus from Carphophis amoenus

 

56

(Say). 2. punctatus is characterized as having a neural

 

Spine which is thicker, higher, and with more of a posterior
overhang than T. amoenus. The fossils resemble Q.

punctatus in these characters.

 

Carphophis amoenus (Say)

 

Material--Four precaudal vertebrae, USNM 209185.
Remarks-~These vertebrae were distinguished from

Diadophis punctatus using the criteria cited under that

 

species.

Coluber or Masticophis

 

Material--Ten precaudal vertebrae, USNM 209186.
Remarks--The vertebrae probably should be assigned

to Coluber constrictor Linnaeus or Masticophis flagellum

 

 

(Shaw), but I am unable to distinguish between the vertebrae

of these two species.

Elaphe, sp. indet.
Material--Eight precaudal vertebrae, USNM 209187.
Remarks--Holman (1965) notes that the vertebral
neural spines of Elaphe vulpina (Baird and Girard) are
lower than those of Elaphe guttata (Linnaeus) and Elaphe
obsoleta (Say). The fossil vertebrae have high neural
spines and thus represent one of the latter two species.

I am unable to separate the vertebrae of those two species.

57

Lampropeltis getulus (Linnaeus)

 

Material--Sixteen precaudal vertebrae, USNM 209188.
Remarks--Auffenberg (1963b) and Holman (1965) have

discussed the vertebral characters of Lampropeltis.

 

Because of their high neural spines, the fossil vertebrae

are assigned to T. getulus.

Lampropeltis triangulum (Lacepede)

 

Material--One hundred seven precaudal vertebrae,
USNM 209189.
Remarks--These Lampropeltis fossils possess low

neural spines and are assigned to T. triangulum. Many of

 

the fossils represent a large individual or individuals,

possibly the northern subspecies T. E. triangulum.

 

Natrix sipedon (Linnaeus)

 

Material-~Seven precaudal vertebrae, USNM 209190.
Remarks--Natrix vertebrae are most easily confused

with Thamnophis vertebrae. Natrix vertebrae are prOpor-

 

tionately wider than Thamnophis vertebrae of equal width.

 

In addition, most Natrix vertebrae possess a prOportionately
higher neural spine. The fossil vertebrae resemble Natrix
in both characters. The fossil vertebrae are referred to

T. sipedon since they possess neural spines which are-
longer than high. This is characteristic of N. sipedon.
Other Natrix species of the southeastern United States

[T. gyclopion floridana Goff, N. erythrggaster Forster,

 

 

N. fasciata Cope, N. rhombifera (Hallowell), and N.

 

58

taxispilota Holbrook] have neural spines which are higher

than long (Holman, 1967).

Storeria dekayi (Holbrook)

 

Material--One precaudal vertebra, USNM 209191.
Remarks--The precaudal vertebrae of the small
natricines are difficult to identify. Auffenberg (1963b)

points out that Storeria differs from Seminatrix in having

 

a longer centrum. He separates Storeria from Haldea by the
ratio of the neural spine length to the zygosphene width,
Storeria showing a ratio greater than 1.21 and Haldea
showing a ratio less than 1.21. The fossil has a value of
1.25 for this ratio, placing it in the genus Storeria. For
the two species of Storeria, Auffenberg lists the neural
spine length to zygosphene width ratio as 1.23 to 1.36 for

S. dekayi and 1.60 to 1.64 for S. occipitomaculta. This

 

would place the fossil in the dekayi group. However, the
ratios quoted are for middle precaudal vertebrae only. If
ratios on all precaudal vertebrae are tabulated there is
considerable overlap of values. Thus the correct identifi-
cation of the vertebrae rests in the ability to distinguish
the middle precaudal vertebrae in disarticulated skeletons.
The fossil vertebra appears to be a middle precaudal

vertebra, and is therefore identified as g. dekayi.

Thamnophis sirtalis (Linnaeus)

 

Material--Six precaudal vertebrae, USNM 209192.

59

Remarks--Thamnophis can be separated from the other
moderate sized natricines of North America by their long,
narrow, vertebral centra (Auffenberg, 1963). Holman (1962b)
separates T. sirtalis from T. sauritus (Linneaus) using the
angle made by the prezngpophyseal processes and the
longitudinal axis of the centrum. Unfortunately, these
processes are missing in the Ladds fossils. Examination of
recent skeletons of ten T. sirtalis and eight T. sauritus
indicates that the neural spine of T. sauritus is lower and
longer than that of T. sirtalis. On that basis the fossils

were identified as T. sirtalis

Family Viperidae
Crotalus horridus Linneaus

Material--Sixty-nine precaudal vertebrae, USNH
209193.

Remarks--Holman (1967) detailed how Crotalus
horridus vertebrae may be distinguished from Crotalus
adamanteus Beauvois, Agkistrodon contortrix (Linneaus),
and Agkistrodon piscivorus (Lacepede). The fossil
vertebrae conform to his diagnosis of g. horridus in
having low neural spines and shallow pits on either side

of the cotyle.

60

DISCUSSION

The Herpetofauna

 

The present study adds Bufo terrestris, Chrysemys

 

 

piggg, and Storeria dekayi to the known Pleistocene
herpetofauna of the Ladds site. The total herpetofauna
now consists of at least twenty-six species.

Species not found in this sample, but previously

reported by Holman (1967), are Desmognathus or Leurognathus,

 

Gyrinophilus sp. indet., Pseudotriton ruber (Sonnini),

 

Hyla crucifer Wied, Anolis carolinensis Voigt, Sceloporus

 

 

undulatus (Latreille), and Heterodon platyrhinos Latreille.

 

Paleoecology

 

The members of the herpetofauna indicate that a
‘moist woodland habitat was present near the deposition

site. Desmognathus, gyrinophilus, and Pseudotriton species

 

are usually associated with a clear woodland stream or
brook (Conant, 1958). The other members of the herpeto-
fauna could all be found in a moist deciduous forest which
contained or bordered an open, brushy area. The presence

of Pseudemys concinna and Clemmys insculpta suggests the

 

forest also contained a large pond or a large body of
running water. Such habitats can be found in the Ladds

area today.

61

Zoogeography and Pleistocene Climate

With the exception of Clemmys insculpta and Bufo

 

terrestris all the species of the Ladds fossil herpeto-

 

fauna can be found living in or near the area today. At

present, the range of g. insculpta extends no further south

 

than northernmost Virginia. B. terrestris is found no

 

further north than mid-Georgia, approximately one hundred
miles south of the Ladds site. These distributions are
shown in figure 8.

Ray (1965), and Lipps and Ray (1967) point out that
there is every Opportunity for fossil mixing at the Ladds
site. Ray (1967) and Holman (1967) indicate that the
occurrence of northern species and southern species in the
same deposit may be indicative of faunal mixing. However,
both acknowledge the possibility that the assemblage could
represent a unit fauna.

Even interpreted as a unit fauna, little climatic
significance can be credited to the sympatric occurrence of

g. insculpta and B. terrestris. The manner in which

 

 

Pleistocene variations in precipitation and temperature may
have affected distributional patterns of various organisms
is poorly known. Not until the physiolOgical and ecological
tolerances of the critical Species are known better, can
definitive faunal-based interpretations of the paleo—

climate be made.

62

 

 

 

 

.9 o a
.ll ’
—7 I
t\
/\
/’ ’
f,” O

 

 

 

 

 

Figure 8. Range limits of Clemmys insculpta and Bufo

terrestris.

----- Northern limits of g. terrestris
----- Southern limits of g. insculpta

 

63

SUMMARY

Three species have been added to the faunal list of
reptiles and amphibians found at the Ladds, Georgia site.
The total recorded herpetofauna now stands at twenty-six
species.

The species represented by the fossils found at
the Ladds site suggest that a moist deciduous forest was
present when the fossils were deposited.

I The herpetofauna gives little indication that the

paleoclimatic conditions were much different from the

climatic conditions that prevail in the area today.

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143 2820

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