BRUCE J. MAC FADDEN

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FLORIDA STATE MUSEUM

Biological Sciences
VOLUME 18 1974 NUMBER 3

CHECKLIST OF FOSSIL LAND TORTOISES
( TESTUDINIDAE)

WALTER AUFFENBERG

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UNIVERSITY OF FLORIDA GAINESVILLE



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CHECKLIST OF FOSSIL LAND TORTOISES (TESTUDINIDAE)

WALTER AUFFENBERGI

SYNOPSIS: This checklist is based on the main literature pertaining to fossil tortoises
and an examination of many important specimens in museums in North and South
America, Europe, and India.2 Zoogeography and evolutionary trends are outlined
and briefly discussed. Of 318 species originally described as fossil land tortoises,
22 are removed from the family and placed elsewhere, 9 are considered nomina nuda,
13 are based on material unidentifiable at the species level. Two primary homonyms
are suppressed: Gophems depressus and Hadrianus robustus, here renamed Copherus
brattstromi and Geochetone gilmorei respectively. Many species formerly placed in
Testudo ( sensu latu) are reallocated to other genera.

i The author is Curator in Herpetology at the Florida State Museum and Professor
of Zoology, University of Florida, Gainesville, Florida 32611. Most of his previous
contributions have been in evolution, behavior, and ecology of land tortoises. He
is currently engaged in research on the Komodo monitor, herpetology of Komodo
Island, Indonesia, and is continuing his tortoise studies. Manuscript accepted 15
July 1971.
2Partially supported by NSF GB 1362 and 2725.

Auffenberg, Walter. 1974. Checklist of fossil land tortoises ( Testudinidae). Bull.
Florida State Mus., Vol. 18, No. 3, pA 121-251.



122 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

TABLE OF CONTENTS

INTRODUCTION _ 122
METHODS AND DEFINITIONS 123
ZOOGEOCRAPHY AND ECOLOGY 124
EVOLUTION, PHYLOGENY, AND TRENDS 127
TAXA INCORRECTLY PLACED IN TESrUDINIDAE _ - 139
NOMINA NUDA AND UNIDENTIFIABLE MATERIAL 140
SYSTEMATICS - - 140

Genus Acinixys 140
Genus Chersina 141
Genus Geochelone 141

Subgenera
Aldabrachelus 142 fHesperotestudo ...... -____ 160
Asterochelys .....___..____._-_._____- 144 Indotestudo 167

t Caudochelys ______...__.__-_.._____ 144 Manouria 168
Chelonoidis 148 iMegalochelys 173

tcylindraspis ....__-________--.-_-._ 150 1 Monachel!/s 174
+Ct,matholcus ..._.-_._-___._____-_- 152 Subgenera Inquirendae 175
Geochelone 153

Genus Copherus - 179
Genus Homopus 187
Genus Kinixys 188
Genus Malacochersus 188
Genus Psammobates 189
Genus Pyxis 189
Genus + Stylemys 190
Genus Testudo 193

Subgenera Pseudotestudo 195
Agrionemys 194 Testudo 195

Genus Uncertain 209
Genera Inquirendae · 211

+Cheirogaster __-__.-_-_-_-__.-_--___-_....__. 211 1 Kanmchelys 212
tfloridemus 212 tSinohadrianus 213

LITERATURE CrrED 214
APPENDIX 236
INDEX TO SPECIES AND SUBSPECIES 247

INTRODUCTION

More than 200 species of fossil and Recent turtles are now assigned
to the family comprising the land tortoises. The fossil species are listed
in the following pages together with the named forms that were at one
time or another considered tortoises. The considerable number of fossil
forms named from material too fragmentary to be determinable have
been placed in  a separate category. Such placement serves a useful pur-
pose in that names of possible, but presently questionable, validity can-
not be treated as having equivalent value with adequately based tax-
onomic categories. Nor can such names be placed in synonymy, as this
action would be only arbitrary and doubtful.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 123

In almost all of even the better known fossil tortoises, the skull re-
mains unknown. In only a few are the limbs and girdles known. Type
specimens are widely scattered in collections, and no worker has ever
seen a majority of both the fossil and Recent forms.

This multiplicity of species and the inadequacy or inaccessibility of
the fossil material make the assessment of phyletic relationships even
more theoretical and tentative than in most other checklists, but it is felt
a complete set of named forms, arranged to suggest relationships that
can be tested, will add something pertinent and perhaps decisive to the
present mass of nonintegrated detail. Accordingly the following is sub-
mitted with a full understanding of its transitory status, because even
with its shortcomings the time seems right for such a summary. It is
hoped that its availability will stimulate contributions, particularly in the
fossil forms that otherwise might not be forthcoming for many years.

METHODS AND DEFINITIONS'

My primary objective has been to list those species of fossil turtles that are now,
or have sometime in the past been considered as belonging to the family Testudinidae.
I have followed no particular rule for including inadequately known fossil species in
this family. In many cases the final criterion was similarity of shape or ornamenta-
tion of various asteological elements between these and known testudinids.

All extant Recent genera are included for convenience. Extant species are listed
only if. they have· also been reported as prehistoric f6ssils.

To make this checklist more useful, the particular set of ground rules to which I
have adhered must be stated:

( 1 ) Use of parentheses: The International Rules of Zoological Nomenclature are
clear on this point ( Article 23) and have been followed throughout. The rule states
that parentheses are to be used only to indicate the changes prescribed,. not indis-
criminately to indicate any change of combination.

( 2) The synonymy includes a reference to the original description or use of each
name. These are usually listed chronologically.

( 3) The nature and location of type specimens, as well as type Iocalities and
horizons, are cited for each species wherever possible.

( 4) Both geographic and geologic ranges are provided for each taxon. In many
instances geologic age is revised from that of the original work in accordance with
more recent standard references on the nomenclature and correlation of continental
fossiliferous deposits.

(5) Our knowledge of fossil tortoises is actually quite meager, so it is impossible
to place each taxon in a definite phyletic position at this time. The arrangement of
all genera, subgenera, and species is alphabetic rather than phyletic.

( 6) The large number of species listed ( 206) may lead some to believe that, I
have recognized an unduly large proportion of species names. In general I have
preserved names for many presently questionable forms until there is some proof that
they represent variant or aberrant individuals. I have taken certain liberties in
synonymyzing Tertiary forms from both Europe and Asia. For the names of Recent
forms I follow mainly Williams ( 1952) and Loveridge and Williams ( 1957). The
question of subspecific relationship for the fossil populations is impossible to deter-
mine on the basis of intergradation, and therefore morphological similarity remains
the sole criterion on which my judgments are based.

A wide possible range in degree of distinctness usually occurs betweell the genera,
subgenera, or species of fossil and Recent forms. The number and kinds of differ-



124 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

ences between fossil species of a single genus that can be accepted as commensurate
with the definition must vary with the investigator, the trend of the times, and.the
nature of the specific group being studied. As the criteria for recognizing full species
in fossils are not yet standardized, it is impossible to be absolutely consistent in the
definition of both fossil and Recent genera, subgenera, or species.

( 7) Every extinct taxon is preceded by the symbol t,
( 8) The present interpretation of the homonym rule requires that the names of

two tortoises are affected as follows:
A. Gopherus idepressus Brattstrom ( 1961), Miocene of California.

Testudo depressa Guerin-Meneville ( 1828 ) ( = Gopherus pol!/Phemus), Re-
cent of southeastern United States.

Gopherus tdepressus Brattstrom is here renamed Gophems tbratestromi in
honor of its discoverer.

B. Hadrianus trobustus Gilmore ( 1915), Uinta Eocene, Utah ( = Geochelone
[Hadrianus] trobusta).

Testudo twbusta Leith-Adams ( 1877), Pleistocene Malta ( = Geochelone
[?Geochelone] frobusta).

Hadrianus trobustus Gilmore is here renamed Geochelone tgilmorei in honor
of its discoverer.

( 9) Species groupings and subgeneric categories of tortoises rest on fairly secure
bases; above this uncertainties increase. Placing almost all of these groups in the
genus Testudo, as is most often done, does not in my opinion refect the apparent
fact that the homogeneity of tortoises is due to similar trends in several phyletic lines.
Therefore the classification of Williams ( 1952) and Loveridge and Williams ( 1957),
rather than the more recent but conservative treatment of Wermuth and Mertens
( 1961 ), is followed.

( 10) Under most generic and species accounts a section headed "Remarks" in-
cludes literature citations to important publications as well as comments on presumed
relationships. No attempt has been made to include every reference to fossil tor-
toises-only those that seem most important for purposes of this checklist.

( 11) Genera not represented by fossils are briefly diagnosed with pertinent re-
marks appended.

( 12) To bring this work to completion no publications were included that ap-
peared after 1 January 1972.

ZOOGEOGRAPHY AND ECOLOGY

The family Emydidae probably has the widest distribution of all the
nonmarine turtle families. The family Testudinidae ( the true tortoises)
has a more restricted distributional pattern. It is here regarded as being
represented by at least 16 genera, 10 of which are still living. Of the
extant genera 5 are Ethiopian endemics ( Psammobates, Manchochersus,
Chersine, Kinixys, and Homopus), 2 are confined to Madagascar ( Aci-
nixys and Pyxis), 1 is widely distributed throughout the southern Pale-
arctic ( Testudo), and 1 is restricted to the southern Nearctic ( Gopherus).
The remaining genus, Geochelone, is the largest, being represented at
least within historic times by about 19 species. It is distributed through-
out much of Africa south of the Sahara, Madagascar, several smaller
islands in the western Indian Ocean, extreme southeastern Asia, including
some of the East Indian islands, extreme southern Central America, most
of South America east of the Andes, some of the Leeward Islands in the



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 125

West Indies, and the Galapagos Islands. Although never native to Aus-
tralia or Melanesia, waif dispersal across marine barriers is common
( Williams 1950a. 1952; Simpson 1942, 1943).

The living tortoises are for the most part subtropical to tropical in dis-
tribution, being most common in subhumid to arid grasslands and savan-
na habitats, though there are a few mesic tropical forest forms. 1 During
the Tertiary their ranges extended throughout what are now temperate
latitudes ( Brattstrom 1961; Hibbard 1960; Auffenberg and Milstead
1965 ) ( Fig. 1). This is believed due to a high degree of climatic equ-
ability, enabling tropical and temperate biotas to intermingle ( Axelrod
1967). As the climate became cooler, the distribution of tortoises was
obviously affected ( Hibbard 1960; Brattstrom 1961). Quaternary tori
toises in the middle and northern latitu(les were subjected to at least four
major periods of colder climate, often more moist. Drier, warmer cli-
mates characterized parts of the three interglacial ages. Each of the
glacial and interglacial periods may have been cooler than the preceding
corresponding periods. Though the evidence is meager, tortoises seem
to have expanded northward during each interglacial. More important,
the northern range limit of tortoises with each successive interglacial was
farther southward. Though the range was smaller during glacial periods,
it seems to have become more restricted with each succeeding advance,
but none of these changes were severe enough to bring about major tor-
toise extinctions. During and after the last glacial, severe drought and
cold in the northern latitudes played important complementary roles in
extinction of large tortoises. Unlike their large contemporaries, smaller
species of tortoises survived these major temperature changes by retreat-
ing into a burrow. It is inconceivable, on mechanical grounds alone that
the extinct giant tortoises of the Pleistocene tunneled. Few living tes-
tudinids of any size burrow in the earth. Those that are known to do so
C Gopherus and some species of Testudo) continue to inhabit the higher
latitudes.

Not all tortoise extinction can be explained solely on late Quaternary
climatic changes. For example it does not fully explain the extinction of
truly gigantic land species in tropical continental areas. Within historic
times such giant species have lived only on islands without large preda-
tors. This has led some workers to suggest that gigantism in tortoises
occurs only in the absence of predation, but this is not so. Each of the

1 Few really complete ecological studies of tortoises are available. The more important ones are:
Beck 1903; Fryer 1911 ; Miller 1932, 1955 ; Hediger 1935 ; Bogert and Cowles 1947 ; Woodbury
and Hardy 1946; Guibe 1950, 1954 ; Cherchi et al. 1958 ; Cernov 1959; Eibel-Eibesfeldt 1959;
Khozatsky 1959; Medem 1960, 1962; Honegger 1964 : Obst and Meusel 1965; Carpenter 1966;
Hutchison et al. 1966; Schmidt-Nielson and Bentley 1966 ; Grubb 1967; Stoddart and Wright
1967 ; Frazier 1968 ; Auffenberg 1969; Auffenberg and Weaver 1969.



126 
B

U
LLE

T
IN

 FLO
R

ID
A
 S

TATE
 M

U
S

E
U

M
 

Vol. 18, N
o. 3

6  --«Me. -/
0

0
O 0 -O 0

0. 0 000

4;. 4.0
o

0

° F

FIGURE 1.-Distribution of land tortoises of the genus Geoche/one, showing reduction of range with time. Diamonds, northern and
southern limits in Oligocene-Miocene. Circles, Phocene-Pleistocene limits. Solid black. and arrows, Recent.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 127

extant elephantine tortoises is represented by gigantic continental rela-
tives. Large carnivores existed in these same continental ·areas from the
Eocene to the Recenti glongside several dominant groups of · these gi-
gantic tortoises. In ·fact the presence of such carnivores may have been
partly responsible for the gigantism·witnessed in several phyletic Iines.

The extinction of giant land tortoises in all parts of the world prob-
ably cannot be explained by a single theory. Man may have played a
role in it, th6ugh only near the clilmination of a keries of dimatic changes
that had already greatly reduced the ranges of the giant species of many
different animals (Webb 1969). Man's repeated visits to those Pacifle
and Indian Ocean islands formerly harboring innum'erable individuals of
gigantic land  tortoises has certainly caused near or complete extinction
in several species. It is also true that in several continental areas the dis-
appearance of some tortoise species can be roughly correlated with the
presence of early man. Two important facts not often mentioned bear
on this problem: (1) s6me very small species of land tortoises. also be-
came, extinct at the same time, and ( 2) the distribution of land tortoises
has.been continuously restricted since the Miocene. The extinction of
the small species cannot be considered part of the general extinction of
large land animals in temperate latitudes that characterized the end of
the Pleistocene. The pre-Pleistocene extinction of tortoise species was
entirely climate-activated. Near the end of the Pleistocene and during
the Early Recent man undoubtedly played a contributory role through
habitat modification, as well as through direct predation on relict popula-
tions.

The only described extinct tortoise genera considered valid in this
contribution. are Floridemys, Cheirogastor, Kansuchelys, Sinohadrianus,
and St!/lemys. Others are either synonyms of presently recognized tor-
toise genera or subgenera or are not testudinids.

-EVOLUTION, PHYLOGENY, AND TRENDS

Extinct species of land tortoises are known from deposits of Middle
Eocene to Recent geologic age. Undoubtedly Paleocene and perhaps
even Upper Cretaceous members of the family will eventually be found.
These prototestudinids will be intermediate between the earliest known
tortoises of the Eocene and primitive members of the Emydidae.

In Eocene deposits of both North America and Africa and the earliest
Oligocene of North America, Asia, and western Europe all the known

1 Lydekker's view that the gigantic tortoises  became extinct in all continental areas at the close
of the Pliocene is now known to be incorrect. Their presence in the Pleistocene is well doc-
umented in North and South America, as well as in Asia, where many gigantic Middle and
Late Pleistocene species are known.



128 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

representatives of the family were in an evolutionary stage represented
by the extant Geochelone emys of Southeast Asia. They were large, low
shelled, plantigrade, nomadic, frugivorous tortoises living in mesic trop-
ical evergreen forests. Much of the evolutionary history of tortoises re-
flects adaptations to grazing in semiarid, subtropical grassland and thorn
forest. The change in habits and ecology is reflected in several major
morphologic evolutionary trends ( Hay 1908, Williams 1950a, 195Ob,
Loveridge and Williams 1957).1

1) The high, convex shell of almost all extant testudinids is evolved
from one that was low and similar to that in Chrysemys. The depressed
shell of a few Recent species of tortoises is probably a specialization re-
lated to shelter utilization. Two reasons have been suggested for the
major trend in changing shell shape: 1) carnivorous land animals cannot
span the greater convexity of a domed shell with their jaws as readily as
a more flattened shell, and 2) the vaulted shell provides greater space
for the lungs. Several workers ( Koch 1934, et aL) have shown that the
capacity of the lungs of testudinids is somewhat greater than that of
equal sized emydids ( except Terrapene, which has a life style similar to
that of testudinids).

2) In emydids the neurals are usually hexagonal, with the broader
end directed anteriorly. The earlier species of tortoises also exhibit this
condition, though in occasional individuals the second neural is octagonal
( sometimes a rare variant in other families of turtlesl ). Among later
fossil testudinids a high degree of neural differentiation is the rule-
usually an alternation of octagonal and tetragonal elements at least an-
teriorly. In some fossil and Recent species the hexagonal neurals have
the broad end directed posteriorly. This is a specialized condition ( Fig.
2). Some extant tortoises retain the emydid condition, except that few
neurals may be tetragonal as in Homopus.

3) Considerable modification has taken place in the pleurals during
the evolution of tortoises. In almost all other turtles the proximal and
distal ends of the second through the sixth pleurals are nearly the same
width. In almost all testudinid species the proximal ends of the second,
fourth, and sixth pleurals are much narrower than the proximal ends of

1 For anatomy of turtles, including tortoises, see Bojanus 1821, Jackson 1837, Parker 1868, Gray
1873c, 1873d, 1873e, Furbinger 1874, Rutimeyer 1874, Zittel 1887-90, Hoffman 1890, Bauer 1891,
Bruhl 1896, Siebenrock 1897, 1898, 1899, 1909, Wieland 1900, Sieglbauer 1909, Willistori 1925,
Beer 1926, Nopsca 1926a, 1931, Ruckes 1929a, 1929b, Bolk et al. 1981-8, Thompson 1932, Vera-
luys 1936. Kuhn 1937, Schepers 1938, 1939, 1948, Zangerl 1939, Walls 1942, Romer 1945, 1956,
Walker 1947, Schumacher 1954, 1955, George and Shah 19558, 1955b, 19598, 1959b, Colby and
Gamble 1957, Parsons 1959, Williams 1959, McDowell 1961, Shah and Patel 1964, Zug 1966,
PawIey 1968.
1 Shell anomalies are discussed by Meyer 1867, Parker 1901, Coker 1910, Lynn 1937, Grant 1946,
Lynn and Ullrich 1950, Mlynarski  1956, Zangerl and Johnson 1957, Staesche 1961, and others.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 129

A 4 <6 <6 <6 <6 <6 <6 <6

8048~ 8 | 4 <6 <6 <6

C 6> 6> 6> 4 <6 <6 <6 <6

D 6> 6> 6> 6> 6> 4 <6 <6

FIcuRE 2.-Generalized neural formula of a pond turtle (A) and land tortoises (B-
D). A. Chrysemys concinna. B. Geochelone tosbomiana. C. Geochelone tcorsoni. D.
Kinixys efosa.

the adjacent elements. In those pleurals that are narrower proximally,
they are expanded distally. Thus the pleural bones appear to be dove-
tailed with one another ( Fig. 3), producing a very rigid dome. The
mechanical advantages of this system have been discussed by Bienz
( 1895). These modifications are generally more conspicuous in later
fossil species within each group. That this differentiation has been
evolved independently in several tortoise genera is suggested by the fact
that the modification takes place early in the fossil record of some genera
and later in others.

4) In most turtles the position of the rib shows distinctly on the in-
ternal surface of each pleural bone, and the rib heads are broad and
thick. The rib heads join their respective centra at the anterior ends of
the latter, and the anterior ribs come into contact with the next anterior
centrum as well. In extant tortoises the ribs are only faintly indicated
on the pleurals, and the rib heads are usually greatly reduced. Early
fossil types tend to have large ribs. Thus there is a gradual reduction in
rib head size with time.

The distal ends of the ribs of tortoises are also reduced when com-
pared with the emydid condition. In most turtles the distal rib ends
project beyond the ends of  the pleural bones, and the projected ends are
usually received in pits in the dorsal edge of the peripherals. In most
extant testudinids the rib ends have all but disappeared, and in adults a



130 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

A -

FIGURE 3.-Pleural differentiation as related  to the neural formula. Alternately wide
and narrow pleurals appear in the mid-Tertiary with the development of octagonal

process of the peripheral rises and enters a notch on the edge of the
pleural ( Fig. 4). The rib ends enter the peripherals only in juveniles.
In early fossil tortoises the emydid character is often retained until the
individual is almost full grown ( Auffenberg 1964d).

5) In most specialized tortoises the epiplastra are thickened for some
distance on either side of the midline and on the dorsal surface. At the
posteri6r border of the thickening the elevation drops off suddenly to the
level of the entoplastron. The ledge thus formed·is more or less deeply
excavated at its base. This excavation and the thickened epiplastra are
not found in the more primitive members of the group ( St!/temys, Homo-



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 131

B

neurals. A. St!/lemys famphithorax, Early Oligocene, Colorado, U.S.A. B. Geochelone
thestema, late Miocene, Colombia, South America.

pus, etc., Fig. 5). When present, the thickened epiplastra usually project
beyond the general curve of the crest of the anterior lobe. This extension
forms the epiplastral lip, which takes several forms in various species and
furnishes valuable specific characters.

6) At the anterior end of the bridge the hyoplastron of each side
sends a process dorsally and anteriorly that articulates with the first
pleural bone. The hypoplastron also sends up an inguinal buttress, which
usually articulates with the anterior half of the lower border of the sixth
pleural. In Stylemys it articulates at the juncture of the fifth and sixth
pleural, as it does in the Emydidae. Thus it seems evident that in land



132 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

A

U4.*nu#'4",M IMUM",1

1
11 4

'6

B

til

f

FIGURE 4.-The primitive pleural rib and peripheral pit articulation of the carapace
of pond turtles ( A, Chrusemys scripta) is modified in most land tortoise groups by
the mid-Tertiary. The pits are generally absent and often replaced by dorsally di-
rected processes ( B, Gopherus po~!/Phemus). The distal end of the pleural  *ibs are
usually weakly developed, or absent.



1974 AUFFENBERC: FOSSIL TORTOISE CHECKLIST 133

A

/<r=3.
f 2-1-.  &:1 ~:-0.~IliijIT- -.,LU --

B

~Emn#+Ut,
f 0 4 0-

1 640 0

0-

0.9
0

-

C
0 0

U 0 .O

e r:.
0 2

00=

FIGURE 5.-Median longitudinal section of the epiplastron. A. Chrysemvs scripta
Pleistocene ( Emydidae). B. Stylemys tamphithorax, Oligocene C Testudinidae). C.
Geochelone tcrassiscutata, PIeistocene ( Testudinidae).

tortoises the articulation tends to be pushed posteriorly with the increased
length of the bridge. The result has apparently been to diminish the
posterior opening of the shell.

7) The shoulder girdle of land tortoises appears to have been modi-
Bed from that of the Emydidae in two respects: the coracoid is greatly
expanded at its medial border, and the procoracoid process makes an
obtuse angle with the body of the scapula. In all Emydidae where the
condition is known, the coracoid is only slightly expanded, and the pro-
coracoid process makes an acute angle with the body of the bone ( Fig.
6).

8) The humerus of the Testudinidae is modified from that of the



134 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

A

-

FIGURE 6.-Shoulder girdle of an emydid, Chrusemus terrapin ( A), and a testudinid,
Geochelone chilensis (B), showing the expanded coracoid of the latter.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 135

A B/ur

CD
I.:*i

t 6

FIGURE 7.-Proximal views of humeri ( A,B) and femora ( C,D) of an emydid,
ChrYsemys terrapin ( A,C) and a testudinid, Geochelone chilensis (B,D).u-ulna
process, r = radial process of humerus.

Emydidae by having both the radial and ulnar processes twisted ven-
trally, with the included angle small ( Fig. 7).

9) Most species of testudinids have five digits on each forelimb, oc-
casionally only four.' The hind foot often has only four digits. No digit
has more than two phalanges ( Fig. 8), emydids have three. The front
foot of Gopherus is unguligrade, the most primitive members tend to be
plantigrade, and advanced members digitigrade.

10) Major trends in the carpus include fusion of subradial elements,
more distal and proximal subulriar elements, and lateral migration of
b6th subulnar and subradial elements accompanying fusion of the me-
dialia ( Fig. 9) ( Auffenberg 1966b ).



136 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

A~ 8/65* 44~ 5 SS 2*4 1~~ 4
~2 93 2

FIGURE 8.-Generalized manus of an emydid, Chrysemys terfapin (A), and a tes-
tudinid, Gopherus polvphemus (B).r-radius, u-ulna. Shaded portions are the
phalanges.

A < Bl <
I r 34 4

 C»

S @ 0 ev 18 8 4 00
FIGURE 9.-Major trends in evolution of the tortoise carpus. A, fusion of subradial
elements. B, fusion of distal and proximal subulnar elements, subradial carpals and
phalanges, as well as the failure of carpal 4 to contact the intermedium (i). C,
lateral migration of subulnar and subradial elements as well as fusion of medialia.
r = radius, u = ulna.

11) The pelvis is constructed on the plan found in the Emydidae.
12) The femur of advanced tortoises is distinguished from that of the

Emydidae by a ridge-like union of the two trochanters. Between this
ridge and the head of the femur there is usually a pit of some depth
( Fig. 7).

13) The most striking differences between the skulls of the Testud-
inidae and those of the Emydidae are found in the excavation of the
former's palate and in the closure of the stapedial notch ( Fig. 10). The



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 137

A

B

l

FIGURE. 10.-The stapedial notch ( arrow) is usually open in emydid turtles ( A,
Chrusemys terrapin) and closed in testudinids ( B, Geochelone chilensis).

palate rises high above the level of the crushing surfaces of the upper
jaws, and the vault thus formed is carried back between the quadrates.
The skulls of most of the earlier fossil tortoises remain unknown.

The division of the family Testudinidae into generic groups is diffi-
cult for a number of reasons: (1) the groups comprising the family are
closely related, and much parallel evolution has occurred, ( 2·) it is diffi-
cult to distinguish between those characters that are merely convergent
and those that may indicate natural divisions, (3) they display various
combinations of primitive characters, (4) advanced characters in the
several lines have developed quite independently, and (5) the geologic
time at which these advanced characters arose differs in various groups.



138 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

The land tortoises undoubtedly arose from primitive emydids, prob-
ably toward the close of the Late Cretaceous. Th@ primitive forms were
probably already well distributed in the Paleocene, but we have no proof
of this. By the Middle Eocene they are already found in all the continen-
tal areas except Australia and South America. The view of Hay ( 1908)
that North America is probably the ancestral home of the family is not
followed here, mainly because tortoises almost as old are now known
from other parts of the world. A considerable degree of evolutionary
differentiation is indicated even in the earliest members of the family,
suggesting a 16ng, completely unknown prefossil history in which any
continent ( other than Australia) might eventually prove to be the an-
cestral home.

The most primitive fossil and living testudinids approach the Emy-
didae in the following characters:

1) Interval between ventral processes of  the prefrontals only moderately widened
2) Temporal arcade strong
3) Prootic well exposed
4) Quadrate not enclosing stapes
5) Anterior neurals hexagonal ( wide end forward)
6) Suprapygal one, anterior'to vertebral-subracaudal sulcus
7) Entoplastron anterior to humeropectoral sulcus
8) No greatly thickened epiplastral projection, and no excavation at its base
9) Carapace not doomed, flattened

10) Rib ends fit info peripheral pits
I 1 ) At least prefrontal and frontal scales present ~
12) Scales on forelimb numerous, not great19 enlarged
13) Femoral tubercles present
14) No tail claw
15) Neither carapace nor plastron hinged
16) Nuchal scute present
17) Vertebrals not greatly convex
18) Carapacial keels weak
19) Supracaudal scute divided
20) Submarginal scute absent
21) Gular scutes faired
22) Anal notch moderate

The most primitive tortoise group is . the subgenus Manouria ( for-
merly Hadrianus), genus Geochelone. Most of the other groups prob-
ably evolved from this group ( Fig. 10). Extant members of the group
are restricted to mesic evergreen forests of southeastern Asia, also the
habitat of the early Tertiary forms. Since that time the major evolu-
tionary changes in both moiphology and behavior in tortoises have been
intimately associated with the development of xeric plant communities
and spreading temperate conditions throughout the world.

Those living testudinids that have been designated.'gigantic' tortoises
all belong to the genus Geochelone. Several extinct genera attained
large, size, which recurs in many groups. ·Some gigantic forms, living and



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 139

extinct, evolved from small species, and some diminutive types evolved
from much larger ones.

TAXA INCORRECrLY PLACED IN THE TESTUDINIDAEl

Testudo tmunda Hay 1920. Pleistocene of Tennessee, U.S.A. ( -Terrapene carolina,
see Auffenberg 1963).

Testudo ·tpromarginata Reinach 1900. Lower Miocene of Frankfort a.m., Germany.
C -Ptychogaster tfmncofurtanus, in part, and T. antiqua, in part, see Glaessner
1935).

Testudo tfeiert;argi Szalai 1930. Aquitanian of Salgotarjan, Hungary. ( -Ptvcho-
gaster f f€iert>aryi, see Glaessner 1933)

Testudo teurgstemum Gervais ( ex-Pomel) 1836. Lower Miocene of France.
C -Ptychogaster teurgstemum, see Bam 1951).

fArchaeochelys pougeti Bergounioux 1938b. ( anything organic?).
f Colossoemus macrococcugeana Rodrigues 1892. Miocene of Peru. ( = Triassic

ammonodont, see Huene 1944).
Testudo flaurea Forster and Becker 1888. ( -Pt!/chogaster tlaurae, see Glaessner

1933).
Testudo tstricklandi Phillips 1871. ( - Protochelys tstricklandi, see Lydekker 1889a).
Testudo fcalarea Fritsch 1893. Upper part of the Lower Miocene of Eger, Bohemia,

Germany. C -Ptuchogaster fcalarea, see Glaessner. 1933).
Testudo 1 Zeithii Sukheswala 1947. E6cene of India. ( = Hydraspis Ze:thii [Carter]

Mukerjee 1949; [ = Carteremys leithii Williams 1958al )
"Testudo" twana Koenig 1825. ( -Puppigems? crassicostatus Owen 1841).
Testudo tangangensis Ping 1930. Archeologically associated in South China. C =Ps«u-

docadia tanyangensis, see Lindholm 1931 and Auffenberg 1962a).
Testudo tminuta Fraas 1870 ( part). Miocene of Steinheim, Germany. C -Clemmys

tsteinheimensis Staesche 1931)
Cinix!/S fcouzieri Bergounioux 1935. Oligocene of France. ( - Ptl,chogaster

tcouzieri).
Emys tgaudini Pictet and Humbert 1856. ( - Kinixys [Ptgchogastet] fgaudini

Portis 1882 [ = Ptychogaste, tgaudini] )
"Testudo" teat)erensis Bravard 1858. ( -Clemmys tbravardi).
Testudo tlamoni (part) Gervais 1859. ( -Clemmgs tuidati)
Hadrianus tallabiatus Cope 1872c. ( -Achilemys tallabiata Hay 1908 =?Emydidae).

Although Kuhn ( 1964) lists Testudo pseudo©indobonensts as appearing on p. 126
of Szalai ( 1934), he is actually referring to Trion!/x pseudouindobonensts Szalai ( p.
134). Kuhn also lists Testudo dumeriliana Lartet ( 1851), which should read Emys
dumeriliana Lartet. His listing of Testudo phopedemontana probably falls in the
same category. However I have not seen the paper in which· this reference was
supposedly made ( Sacco 1889).

1Wermuth (1956) discusses the very broad 18th Century use of the generic term '*Testudo" for
nontestudinid turtles. Names involved are Testudo boddaerta ( =Trionvz cartilaginosis), Testudo
brevi-caudata (=Terrapene c. carolinG), Testudo caouana ( =Caretta caretta), Testudo care-
liniang ( =Terrapelle carolina), Testudo dersata ( =Geoemyda punctularia), Testudo Ambria
( =Chelus jimbriatus), Testudo flava ( =Emys btanding£), Testudo granulosa (=Triont,z punc-
tatus), Testudo marina vldgaris ( =Cheionid mydas), Tebtudo meteagris ( =Emis blandingi),
Testudo membranacea ( = ?Triont,z cartilaginosus), Testudo mydas minor ( =Lepidocheljs o.
kempi), Testudo nasicorms ( =Caretta caretta), Testudo pianitia ( =Macroclemys temminki),
Teetudo punctata (=Emys orbicularis), Testudo rubicunda ( =Pelomedusa submfa), Testudo
mgosa (=Chelonia m. japonica), Testudo semimembranacea ( =Trionvz sinens€s), Testudo ser-
pentina (=Chelydra serpentina), Testudo striata (=Trionl/z cartitaginogus and T. tnunglds
part), Testudo vernicosa (=Geoemvda pulictularia), Testudo viridi-squamosa (=Lepidochell/8 0.
kempi).



140 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

NOMINA NUDA AND UNIDENTIFIABLE MATERIAL

The following list includes those names I consider lacking a proper diagnosis.
Names in parentheses indicate authors who have previously taken the same position.
Testudo tminuta Bravard 1844. ( Szalai 1934).
Testudo imedia Bravard 1844. ( Staesche 1931).
Testudo twalensis Khozatsky 1945.
( Testudo) thouzei Dollo 1912. Only the species name was used originally and later

assumed to be Testudo by Bergounioux ( 1933b).
Testudo tnumurensis Meyer 1865. ( Kuhn 1964).
Testudo taustralls Moreno 1889. ( Williams 1950a).
Testudo trisgoviensis Fraas 1870. ( provisional name).
Testudo tformosa Moreno 1889. ( Williams 195Oa).
Testudo tparanensis Scalabrini 1884.

Names based on material here considered unidentifiable at the species level in-
clude the following forms. Authors who have previously suggested such actions are
in parentheses.
?Testudo tnerandi Gray 183la.
Testudo + cuvieri Fitzinger 1835, ( = Testudo radiata fossilis Meyer 1832; = Testudo

radiata Pictet 1845).
Machrochelys tmira Meyer 1858, ( = Testudo [Macrochelys] mira Zittel 1889).
Stylemys toregonensis Leidy 1871c. ( = Testudo toregonensis Leidy 1873; =

?Stylemys tnebrascensis Cope 1885),
Testudo tfrizaciana Lartet 1851 ( = Stylemys f frizaciana Auffenberg 1964d) ( Pictet

1853).
Testudo tdespotti Szalai 1934. ( Kuhn 1964).
Testudo thungarica Szalai 1934.
Testudo flambrechti Szalai 1934.
Testudo tstrandi Szalai 1936. ( Mlynarski 1966a).
Testudo tgigas Bravard 1844.
Testudo tlemanensis Bravard 1844. ( Riabinin 1926).
Testudo tracmecskeensis Szalai 1932 ( Ms. name), 1934. ( Glaessner 1935 ).
Testudo tpygmaea Lartet 1851, ( = St!/lemys 1 pugmaea Auffenberg 1964d) ( Maack

1869).

SYSTEMATICS
Class REPTILIA Laurent 1768, p. 23.
Order TESTUDINATA Shaw 1802, p. 5.
Suborder CRYPTODIRA Cope ( part) 1870, p. 123.
Family TESTUDINIDAE Gray 1825, p. 210.

DEFINITION.-Cryptodiran turtles with no more than two phalanges
in the digits of either front or hind feet; carapace usually high-arched;
stapes always encloked by quadrate; surangular developed on outer sur-
face of jaw; splenial absent; dorsal rib heads tending to be vestigial; pubis
joins ischium on same side below fenestra.

Genus Acinixys Siebenrock
Testudo Grandidier 1867, p. 233 ( part).
Puxis Boulenger 1889, p. 145 ( part).
Acinixys Siebenrock 1903b, p. 244-6, pl. 33-34.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 141

GENOTYPE.-Acinixys planicauda ( Grandidier).
DEFINITION.-A monotypic Ethiopian genus, restricted to the Mal-

agasy Republic and characterized by contours of the rounded anal scales,
which are not divided by a median notch.

GEOLOGIC RANGE.-No fossils known.
GEOGRAPHIC RANGE.-Malagasy Republic.
REMARKS.-Unfortunately a poorly known genus deserving consider-

able study and attention.

Genus Chersina Gray
Testudo Thunberg 1795, p. 3 ( part).
Chersine Merrem 1820, p. 38 (not of Linnaeus).
Chersina Gray 183la, p. 7, 14 ( not of Humphreys).
Goniochenus Lindholm 1929, p. 285.
Neotestudo Hewitt-1931 , p . 504.

GENOTYPE.-Testudo angulata Schweigger ( = Chersina angulata
[Schweigger]) by monotypy.

DEFINITION.-A monotypic endemic Ethiopian tortoise characterized
by having no hinges on either the carapace or plastron and a single gular
scute on a strongly-projecting and greatly thickened epiplastral projec-
tion.

GEOLOGIC RANGE.-No fossils known.
GEOGRAPHIC RANGE.-Cape of Good Hope Province, South Africa.
REMARKS.-Rarity of double gulars in this species ( Cairncross 1958,

Archer 1961) may be of importance in future finds of fossil relatives.

Genus Geochelone Fitzinger
Chenine Merrem 1820, p. 29 ( part).
Testudinites Weiss 1830, p. 293.
Geochetone Fitzinger 1835, p. I 11.
Megalochelys Falconer and Cautley 1837, p. 358 (part ).
Colossochelys Falconer and Cautley 1844, p. 54.
Geoem{/da Cantor 1847, p. 2 ( part).
Teleopus Le Conte 1854, p. 187 ( T. luratus)
Manouna Gray 1854, p. 134.
Emys Milne-Edwards In Grandidier (part) 1868, p. 1167.
Scapia Gray 1869, p. 169 ( S. falconeri).
Stglemys Cope 1870, p. 124 ( part).
Peltastes Gray 187Ob, p. 655 ( part).
Gopher Gray 1870a, p. 190.
Testudo Leidy 187la, p. 154 ( part).
Centrochelys Gray 1872a, p. 5 ( T. sulcata Miller, by monotypy).
Hadrianus Cope 1872a, p. 2 ( part).
Stigmochelus Gray 1873a, p. 5 ( T. pardatis Bell, by monotypy).
Elephantopus Gray 1873b, p. 724 (part).
Eupachemys Leidy 1877, p. 7 ( nomen nudum).
Homopus Boettger 1893, p. 8 ( part).



142 BULLETIN FLORIDA STATE MUSEUM ' Vol. 18, No. 1

Pampatestudo Lindholm 1929, p. 285 ( part).
Megachersine Hewitt 1931, p. 257 ( T. pardalis Bell, by original designation).
Gopherus Williams 195Oa, p. 30 ( part).

GENOTYPE.-Testudo steNata. Schweigger ( - T. elegans Schoepff ).
DEFINITION.-An almost cosmopolitan tortoise genus with triturating

surface of maxilla strongly ridged; median premaxillary ridge absent;
maxillary not entering roof of palate; anterior palatine foramina small,
concealed in ventral view; prootic typically well exposed dorsally and
anteriorly; quadrate usually enclosing stapes; subangular subequal in
height to prearticular; neck with second, third or fourth centrum bi-
convex.

Carapace never hinged; typically the anterior neurals alternately
octagonal and quadrilateral; outer side of third costal scute about as long
as, or longer than that of the fourth; no submarginal scute; two suprapy-
gals, the anterior larger, bifurcating posteriorly to embrace the smaller
posterior elements, which ( in post-Eocene forms) is crossed near its
middle by the Sulcus between the fifth vertebral and the supracaudal.

Plastron not hinged; gular region more or less thickened and pro-
duced; gulars single or paired, longer than broad.

GEOLOGIC RANGE.-Eocene to Pleistocene of North America, Eocene
to Pliocene of Europe, Oligocene to Recent of Asia, Oligocene and Mio-
cene of Africa, Miocene to Recent of South America, and Pleistocene to
Recent of the West Indies and a number of islands in the Indian Ocean.

GEOGRAPHIC RANGE.-GalapagOS Islands, South America, West Indies,

Africa, Malagasy Republic, islands of the Indian Ocean, Southern Asia,
East Indies, and Ceylon.

REMARKS.-A combination of primitive and advanced characters is
common in species of this genus. This large tropical, nearly cosmopoli-
tan genus contains the large extant mainland and insular tortoises as well
as all the extinct giant tortoises. It is divided into the following 13 sub-
genera.

Subgenus Aldabrachel!/s Loveridge and Williams

Aldabmchelus Loveridge and Williams 1954, p. 225 (as subgenus).
Testudo Schweigger . 1812, p . 327 .
Emys Milne-Edwards In Grandidier 1868, p. 1167.

TYPE SPECIES.-Testudo gigantea Schweigger.
DEFINNION.- A subdivision of the genus Geochelone known only

from the Aldabra Islands and the Malagasy Republic. The nuchal scute
is present or absent; first dorsal vertebra short; gular scutes paired but
not divergent; entoplastron not crossed by the humero-pectoral sulcus;
external nares higher than wide; quadrate enclosing stapes or not.

GEOLOGIC RANGE.-Pleistocene ( of Malagasy Republic) to Recent.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 143

GEOGRAPHIC RANGE.-Aldabra Islands, Seychelles Archipelago, and
Malagasy Republic, but probably more widely distributed in the past
( Agalega, Assumption, Amirante, Astove, Alphonse,'Africa, Providence,
St. Pierre, Farquhar, Cosmoledo, Gloriosa, and · possibly even Chagos
Islands [Rothschild 1915, Fryer 1911 ]).

REMARKS.-For the best monographic treatment of the extant species
( G. elephantina), see Rothschild ( 1915). The several fossil specimens
from Europe referred to this group under Testudo elephantina of Dum-
eril and Bibron ( Pictet 1845, Pomel 1846, Maack 1869, and Bergounioux
1938a) obviously belong to the subgenus Geochelone. Testudo tgigantea
Bravard 1844 ( not of Schweigger) is a nomen nudum. Only one species,
G. gigantea Schweigger 1812, is extant. It is confined to the Aldabra
Islands, Seychelles Archipelago, and in the Indian Ocean. For skeletal
morphology see Giinther ( 1877a, 1877b) and Loveridge and Williams
( 1957). At least three species and one subspecies are extinct.

Geochelone ( ?Aldabrachelys) fabrupta C Vaillant)

Testudo abrupta Milne-Edwards In Grandidier 1868, p. 1'161 ( nomen nudum).
Testudo abrupta Vaillant 1885b, p. 874.
Testudo grandidier Boulenger 1894, p. 305 (part).

TypE.-Museum of Natural History ( Paris); carapace and limb
bones.

TypE LOCALI'rY AND HORIZON.-Amboulitsate in central part of the
Malagasy Republic; Late Pleistocene.

GEOLOGIC RANGE.-Pleistocene only (?).
GEOGRAPHIC RANGE.-Central Malagasy Republic.
REMARKS.-The subgeneric affinities of this species are not clear.

Geochelone (Aldabrachelys) gigantea f gou#ei ( Rothschild)
Testudo gou#ei Rothschild 1906, p. 753 ( ?Farquhar Island).

TYPE.-Tring Museum; a mounted adult specimen.
TYPE LOCALITY.-?Farquhar Island, Seychelles Archipelago, Indian

Ocean.
GEOLOGIC RANGE.-Recent, but now extinct.
GEOGRAPHIC RANGE.-Known from a single specimen of uncertain

origin.
REMARKS.-The type appears to be a unique form. Skeletal mor-

phology of the extant subspecies is discussed by Gunther ( 1877a, 1877b).

Geochelone ( Aldabrachelys) fiumeirei C Sauzier)
Testudo sumeirei Sauzier 1893, p. 7 ( Mauritius?, in error).
Geochelone ( Aldabrachelys) sumeirei Loveridge and Williams 1957, p. 225.



144 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

TYPE.-Tring Museum; a mounted adult.
TYPE LocALITY.-Seychelles Islands?
GEOLOGIC RANGE.-Recent, but· now extinct.
GEOGRAPHIC RANGE.-Seychelles Archipelago?, Indian Ocean.
REMARKS.-A note on the type states that it is one of Bve specimens

taken from the Seychelles to Mauritius by Chevalier Marion de Fresne in
1766 as a gift for the Port Louis Garrison. This is the famous "Marion's"
tortoise, which died accidentally after 152 years in captivity.

Geochelone ( Aldabrachelys) tgrandidieri ( Vaillant)
Emys gigantea Milne-Edwards In Grandidier 1868, p. 1165 (not of Schweigger

preoccupied).
Testudo grandidieri Vaillant 1885b, p. 874.
Testudo gigantea Boulenger 1892, p. 581 ( part).

TypE.-Museum of Natural History ( Paris); parts of shell.
TYPE LOCALITY AND HoRIZON.-Northern Malagasy Republic; late

Pleistocene and/or Recent.
GEOLOGIC RANGE.-Late Pleistocene and ?Recent.
GEOGRAPHIC RANGE.-Northern Malagasy Republic.

REFERENCES.-Skeleton: Boulenger 1894, Auffenberg 1966b.

Subgenus Asterochelys Gray
Asterochel!/s Gray 1873a, p. 4.
Testudo Vaillant 1885a, p. 440.
Geochelone Loveridge and Williams 1957, p. 342.

TYPE SPEcms.-Testudo yniphora Vaillant ( = Geochelone [Astero-
chelys] yniphora [Vaillant]).

DEFINITION.-A subgenus of the genus Geochelone known only by
two living species from the Malagasy Republic. Characterized by pres-
ence of a nuchal scute, gulars single or double, and external nares not
expanded vertically.

GEOLOGIC RANGE.-NO fossils known.
GEOGRAPHIC RANGE.-Malagasy Republic.
REMARKS.-The two presently recognized extant populations ( G.

radiata and G. yniphora) are probably conspecific. For skeletal mor-
phology see William 195Ob, Vaillant 1905, Siebenrock 1897.

Subgenus t€audochelys Auffenberg

Caudochdys Auffenberg 1963, p. 69 ( as subgenus).

TYPE SPECIES.-Testudo crassiscutata Leidy ( = Geochelone [Caudo-
chelys] crassiscutata [Leidy] ).

DEFINITION.-Extinct Nearctic subdivision of genus Geochelone with



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 145

narrow nuchal scute; entoplastron about as wide as long; pectoral scutes
reduced along midline; limbs and tail heavily armored with dermal os-
sicles; above tail, ossicles never fuse to form supracaudal buckler; caudal
vertebrae normal, not compressed or fused, without greatly elongated
transverse processes.

GEOLOGIC RANGE.-Miocene to end of Pleistocene.
GEOGRAPHIC RANGE.-Central and eastern North America.
REMARKS.-Mlynarski ( 1969a, p. 87) mistakenly confuses this sub-

genus with the subgenus Hesperotestudo. Caudochelys is presently con-
sidered as being comprised of at least nine extinct species.

Geochelone ( Caudochelys) tannae ( Hay)
Testudo crassiscutata ? Hay 1916b, p. 11, pls. 1-3 (part).
Testudo annae Hay 1923, p. 114.
Geochelone annae Auffenberg 1963, p. 94.

TypE.-Dr. Mark Francis collection, Texas A&M Univ.; right
epiplastron, anterior part of carapace.

TypE LoCALITY AND HORIZON.-Brazos River at Pittbridge, Burleson
County, Texas, U.S.A.; Aftonian faunal age; Early Pleistocene.

GEOLOGIC RANGE.-Pleistocene.
GEOGRAPHIC RANGE.-Texas, U.S.A.
REMARKS.-This probably is a synonym of G. crassiscutata.

Geochelone ( ?Caudochelys) tbrontops ( Marsh)
Testudo brontops Marsh 1890, p. 179, pl. 8.
Geochelone brontops Auffenberg 1963, p. 87.

TYPE.-Yale Peabody Museum; a shell.
TYPE LOCALITY AND HORIZON.-S. E. corner of Pennington County,

South Dakota, U.S.A.; "Titanother beds" of Indian Creek, Chadronian
faunal age, Early Oligocene.

GEOLOGIC RANGE.-Early Oligocene.
GEOGRAPHIC RANGE.-South Dakota, U.S.A.

Geochelone ( Caudochelys) tcrassiscutata C Leidy)
Eupachemys obtusa Leidy 1877, p. 232.
Eupachemus rugosus Leidy 1889, p. 29 ( error).
Testudo crassiscutata Leidy 1889, p. 31.
Testudo obtusa Hay 1908, p. 458.
Testudo ocalana Hay 1916a, p. 45.
Testudo distans Hay, 1916a, p. 48.
Testudo sellardsi Hay 1916a, p. 49.
Testudo luciae Hay 1916a, p. 52.
Gophems ocalana Williams 195Oa, p. 30.
Geochelone distans Ray 1957, p. 126.



146 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

Geochelone luciae Ray 1957, p. 126.
Geochelone seUardsi Ray 1957, p. 126.
Geochelone ( Caudochelys) crassiscutata Auffenberg 1963, p. 70.

TYPE.-U.S. National Museum; parts of plastron, femur and tibia.
TypE LOCALIT¥ AND HORIZON.-Shoals of Peach Creek, near Arcadia,

DeSoto County, Florida, U.S.A.; Rancholabrean mammalian faunal age,
Late Pleistocene.

GEOLOGIC RANGE.-Middle to Late Pleistocene,
GEOGRAPHIC RANGE.-Florida, north to South Carolina ( Auffenberg

1963), west to at least eastern Texas, U.S.A. ( Holman 1969).
REMARKS.-For best description see Loomis ( 1927) and Auffenberg

( 1963). Probably much more widely distributed than present records
indicate.

Geochelone ( Caudochelys ) f ducatelli ( Collihs and Lynn )
Testudo duet:telli Collins and Lynn 1936, p. 166, pls. 3-4.
Testudo' ducateli Kuhn 1964, p. 116 ( typographical error).
Geochelone duet*elli Auffenberg 1964a, p. 3.

TYPE.-U.S. National Museum; plastron and parts of a carapace.
TYPE LOCALITY AND HORIZON.-3.4 miles south old Chesapeake

Beach RR Station, Maryland, U.S.A.; Zone 10, Calvert Formation, Barsto-
vian faunal age, late Miocene.

GEOLOGIC RANGE.-L~te Miocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-Closely related to the Texas Miocene Geochelone wil-

liamsi ( Auffenberg 1964a).

Geochelone ( ?Caudochelys) ffrancisi ( Hay)
Testudo francisi Hay 1923, p. 116, pl. 8.

TypE.-Dr. Mark Francis collection, Texas A&M Univ.; an epiplas-
tron.

TypE LoCALITY AND HORIZON.-Temple, Bell County, Texas, U.S.A.;
referred to Aftonian faunal age, Early Pleistocene by Hay ( 1923).

GEOLOGIC RANGE.-Pleistocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-This species is probably close to G. crassiscutata

Geochelone ( Caudochelus) thayi C Sellards)
Testudo hal/i Sellards 1916, p. 235, Bg. 7,9
Testudo louisekressmani Wark 1929, p. 401.
Gopherus ha!/i Williams 195Oa, p. 30.
Geochelone ( Caudochdp) ha!/i Auffenberg 1963, p. 78.
Geochelone louisekressmani Auffenberg 1963, p. 79.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 147

TYPE.-U.S. National Museum; a partial shell.
TYPE LOCALITY AND HORIZON.-Near Nichols, Polk County, Florida,

U.S.A.; Bone Valley Gravel Formation, Hemphillian faunal age, Middle
Pliocene.

GEOLOGIC RANGE.-Middle Pliocene.
GEOGRAPHIC RANGE.-Central to northern Florida.
REMARKS.-C~OSe and perhaps ancestral to the Blancah G. campester.

Geochelone- ( ?Caudochelys) tmilleri ( Brattstrom)
Testudo milleri Brattstrom 1961, p. 546, figs. 5-6.

TypE.-Univ. of California Museum of Paleontology; a partial shell.
TypE LOCALITY AND HoRIzoN.-Barstow syncline, Mojave Desert,

San Bernardine County, California, U.S.A.; Barstow beds,  Barstovian
faunal age, Late Miocene.

GEOLOGIC' RANGE.-Late Miocene.
GEOGRAPHIC RANGE.-Southern California, U.S.A.
REMARKS.-According to Brattstrom ( 1961) it is close to Geochelone

tedwhitei, but this is not certain.

Geochelone ( Caudochelys) ttedwhitei C Williams)
Testudo tedwhitei Williams 1953b, p. 537, figs. 1-3.
Geochelone ( Caudochelys) tedwhitei Auffenberg 1963, p. 80.

TypE.-Museum of Comparative Zoology; a complete plastron.
TYPE LOCALITY AND HORIZON.-Thomas Farm, Gilchrist County,

Florida, U.S.A.; Hawthorne Formation, Hemingfordian faunal age, Mid-
dle Miocene.

GEOLOGIC RANGE.-Middle Miocene.
GEOGRAPHIC RANGE.-Now known from several localities in northern

Florida, U.S.A.

Geochelone ( Caudochelys) twilliamsi Auffenberg
Geochelone willian:si Auffenberg 1964a, p. 3, figs. 1-2.

TypE.-Holotype, Univ. of Texas-Bureau of Economic Geology; a
complete shell.

TypE LoCALITY AND HORIZON.-Garvin Gully, 2 mi. north of Navasota,
Grimes County, Texas, U.S.A.; Garvin Gully local fauna, lower Oakville
Member, Oakville Formation, Arikareean faunal age, Early Miocene.

GEOLOGIC RANGE.-Early Miocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-Presumably close to G. ducatelli ( Auffenberg 1964a).



148 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

Subgenus Chelonoidis Fitzinger
Testudinites Weiss 1830, p. 293.
Chelonoidis Fitzinger 1856, p. 112.
Gopher Gray 187Oa, p. 190.
Elephantopus Gray 1873b, p. 724 ( part).
Pampatestudo Lindholm 1929, p. 285 ( part).

TYPE SPECIES.-Testudo denticulata Linnaeus ( = Geochelone [Chelo-
noidis] denticulata [Linnaeus] ).

DEFINITION.-A Neotropical division of the genus Geochelone with-
out a nuchal scute, entoplastron large, and horny shields with little or
no trace of radiating pattern.

GEOLOGIC RANGE.-Miocene to Recent,
GEOGRAPHIC RANGE.-Recent of southeastern Panama, south over

most of South America east of the Andes to south central Argentina, the
Galapagos Islands, and a number of islands in the Caribbean Sea.

REMARKS.-For best description see Williams ( 195Oa). The sub-
genus is comprised of four extant and six extinct species. The extant
species are Geochelone denticulata, G. carbonaria, G. elephantopus, and
G. chilensis, of which only G. chilensis is known as a fossil.

REFERENCES.-Skeleton: Gray 1855, Fritsch 1871, Gunther 1875, 1877a, 1896,
Jeude 1896, Siebenrock 1897, Heller 1903, Ruckes 1937, Williams 195Oa, Zangerl
1957, Auffenberg 1966b, 1971. Zoogeography: Bauer 1889, Simpson 1943, Auffen-
berg 1971.

Geochelone (Chelonoidis) chile·nsis (Gray)

Testudo sulcata Dumeril and Bibron 1835, p. 74 ( part).
Testudo sulcata d'Orbigny 1847, p. 6 ( not of Gmelin).
Testudo (Gopher ) chilensis Gray 187Oa, p. 190 ( Chile, in error).
Testudo argentina Sclater 1870, p. 471 ( substitute name for T. chilensis Gray

187Oa).
Geochelone chilensis Williams 1960, 8 10.

TypE.-British Museum ( Natural History); two mounted specimens.
TypE LoCALITY.-Near Mendoza, Argentina.
GEOLOGIC RANGE.-Certain fossils from Pleistocene deposits in Ar-

gentina are assignable to this species ( Auffenberg 1971).
GEOGRAPHIC RANGE.-Southwestern Bolivia, Western Paraguay; and

Western Argentina south to near 400 latitude.
REFERENCES.-Skeleton: Auffenberg 1966b, 1971.

Geochelone ( Chelonoidis) tcubensis ( Leidy)

Testudo cubensis Leidy 1868, p. 179.
Geochelone cubensis Auffenberg 1967, p. 37.

TypE.-Philadelphia Academy of Natural Sciences; part of a first
right pleural.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 149

TypE LOCALITY AND H0RIZON.-Chapepote Springs, Banas de Ciego
Montero, Las Villas Province ( = Santa Clara Province), Cuba; Late
( ?) Pleistocene.

GEOLOGIC RANGE.-Late Pleistocene.
GEOGRAPHIC IUNGE.-Brazil.
REMARKS.-Incorrectly stated as Pliocene by a few earlier workers.

Geochelone (Chelonoidis) felata (Gervais)
Testudo elata Gervais 1877, p. 283, pl. 7.

TypE.-Museum of Natural History ( Paris); fragments of dentary,
limbs, and plastron.

TYPE LOCALrrY AND HORIZON.- Lower region, Amazonian basin,"
Brazil; Pampean faunal age, Late Pleistocene.

GEOLOGIC RANGE.-Known only from Late (?) Pleistocene deposits.
GEOGRAPHIC RANGE.-Central to eastern Cuba.
REMARKS.-See WilliamS ( 1952) for best description.

A giant species, probably a synonym of G. sellowi.

Geochelone (?Chelonoidis) tgallardoi ( Roverto)
Testudo gallardoi Roverto 1914, p. 115.
Testudo praestans Roverto 1914, p. 176.

TypE.-Museo Nacional Historie Naturele ( Buenos Aires ); a ·fairly
complete shell.

TypE LOCALrrY AND HORIzoN.-Catamarca, Argentina; Araucanian
faunal age.

GEOLOGIC RANGE.-Late Pliocene.
GEOGRAPHIC RANGE.-Argentina.
REMARKS.-Study of the type specimens of both gallardoi and

praestans has convinced me they are conspecific.

Geochelone (Chelonoidis) tgringorum (Simpson)
Testudo gringomm Simpson 1942, p. 1, Bgs. 1-2.

TypE.-American Museum of Natural ~ History; a plastron and most
of a carapace.

TypE LOCALITY AND HoR,zoN.-South side of Chubut Valley, between
Gaiman and Dolavon, Chubut Territory, Argentina; High in Patagonian
section overlying the Angosturas laminated beds, Miocene ( probably
Early Miocene, according to Simpson 1942).

GEOLOGIC RANGE.-Miocene ( Early?).
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-Ancestral to Geochelone chilensis C Auffenberg 1971).



150 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

Geochelone (Chelonoidis) hesterna Auffenberg
Geochelone hesterna Auffenberg 1971, p. 106.

TypE.-Univ. of California Museum of Paleontology; a complete
shell and partial skeleton.

TYPE LOCALITY AN'b HORIzoN.--3 km northeast of Villavieja, Huila,
Colombia, South America; Hondo group, Cerbatana gravels and clays, La
Venta fauna, Late Miocene.

GEOLOGIC RANGE.-Late Miocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-Intermediate between G. carbonaria and G. denticulata,

though closer to the latter ( Auffenberg 1971).

Geochelone ( Chelonoidis) fsellowi ( Weiss)
Testudinites sellovii Weiss 1830, p. 293.
Testudo seltooli Giebel 1847, p. 53.
Testudo sellowi Couto 1948, p. 1, pl. 1.

TypE.-Museum of Humbolt Univ. ( Berlin ); 8 pieces of carapace
and plastron.

TypE LoCALITY AND HORIZON.-Rio Quequay, Paysandu, Uruguay;
Pleistocene.

GEOLOGIC RANGE.-Pleistocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-Briefly redescribed by Couto ( 1948), who also repub-

lished some of Weiss' · original plates.

Geochelone ( Chelonoidis) tsombrerensis ( Leidy)

Emys sombrerends Leidy 1868, p. 180.
Testudo sombrerensis Williams 1952, p. 552.
Geochelone sombrerensis Auffenberg 1967, p. 35.

TypE.-Academy of Natural Sciences ( Philadelphia ) ( Lost ?).
TypE LoCALIFY AND HORIZON.-Sombrero Guano, Sombrero Island,

West Indies; ? Late Pleistocene.
GEOLOGIC RANGE.-Late Pleistocene.
GEOGRAPHIC RANGE.-Type locality.

REMARKS.-For the most recent description and comparisons see

Auffenberg ( 1967).

Subgenus fCylindraspis Fitzinger

Cylind,aspis Fitzinger 1835, p. 112.

TYPE SPECIES.-Testudo indica uosmaeri Shaw C -Geochelone [Cyl-

indraspis] uosmaeri [Shaw] ).



1974 . AUFFENBERC: FOSSIL TORTOISE CHECKLIST 151

DEFINmON.-An extinct subgenus of the genus Geochelone repre-
sented by four species restricted to the islands of the Mascarene group
in the Indian Ocean. They are characterized by the absence of anuchal
scute, undivided supracaudaI, and single gular.

GEOLOGIC RANGE.-Late Pleistocene to Recent, now extinct.
GEOGRAPHIC RANGE.-Rodriguez, Reunion, and Mauritius  Islands,

Indian Ocean.
REMARKS.-A very poorly known group deserving study with a mod-

ern systematic approach. Giinther ( 1877a) gives a very good resume of
the skeletal characters of many of the named forms.

Geochelone ( Cylindraspis) f indica ( Schneider)
Testudo indica Schheider 1784, p. 355.
Testudo indica permultii Shaw 1802, p. 25.
Chersine fetusa Merrem 1820, p. 29,
Testudo perraultii Dumeril and Bibron 1835, p. 126.

TypE.-Location unknown to author; shell.
TypE LoCALITY.-Reunion Islands, Mascarene Group, Indian Ocean.
GEOLOGIC RANGE.-Recent, now extinct.
GEOGRAPHIC RANGE.-Reunion Islands, Indian Ocean.

Geochelone ( Cylindramis) tgrayi ( Dumeril and Bibron)
Testudo graii Dumeril and Bibron 1835, p. 155.
Testudo inepta Gunther 1873, p. 397 ( Mauritius).
Testudo triserrata Gunther 1873, p. 397 ( Mauritius).
Testudo grayi Giinther 1877a, p. 43.
Testudo leptocnemis Giinther 1877a, p. 47 ( Mauritius).
Testudo microtympanum Boulenger 1890c, p. 4 ( Mauritius?).
Testudo sauzie,i Gadow 1894, p. 315 ( Mauritius).

TypE.-Museum bf Natural History ( Paris ); a shell.
TypE LOCALITY.-Unknown, presumably Mauritius Island, Indian

Ocean ( Gunther 1877a).
GEOLOGIC RANGE.-Late Pleistocene to early 19th Century.
GEOGRAPHIC RANGE.-Mauritius Island.
REMARKS.-All the named Mauritius forms are here placed in the

same species. Gunther ( 1877a) provides an excellent resume of the
skeleton. The data available show that ail insular tortoise populations
are more variable than those of the mainland. This greater variation
has led to considerable taxonomic confusion regarding tortoise popula-
tions in the Mascarene, Aldabra, and Galapagos Islands.

Geochelone ( Cylindraspis) tpeltastes ( Dumeril and Bibron)
Testudo peltastes Dumeril and Bibron 1835, p. 138.
Testudo vosmaeri Fitzinger 1826, p. 1 ( part).



152 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

TypE.-Museum of Natural History ( Paris); dried specimen, less
head and tail.

TypE LOCALTTY,-Rodriguez Island, Mascarene Group, Indian Ocean.
GEOLOGIC RANGE.-Recent, now extinct.

GEOGRAPHIC RANGE.-Rodriguez Island, Indian Ocean.
REMARKS.-Gunther (1877a) and Boulenger (1890c) considered the

specimens of peltastes to be the young of cosmaeri. The presence of an
adult female peltastes and Rothschild's ( 1915, p. 441) insistence on the
ankylosis of the small shells in the British Museum leads me to recognize
G. peltastes as a distinct species for the present.

Geochelone ( Cylindraspis) foosmaeri ( Shaw)
Testudo indica Schoepff 1792, p. 103. (part).
Testudo indica cosmaeri Shaw 1802, p. 27.
Testudo ©osmaeri Fitzinger 1826, p. 44.
Testudo rodericensts Gunther 1873, p. 397 ( preliminary note).
Testudo boutonii Gunther 1875, p. 43.
Testudo col'nTnersoni Vaillant 1889, p. 134.

TypE.-British Museum ( Natural History); shells, skull, and bones.
TypE LOCALITY.-Rodriguez Island, Mascarene Group, Indian Ocean.
GEOLOGIC RANGE.-Recent, now extinct.
GEOGRAPHIC RANGE.-Rodriguez Island, Indian Ocean.
REFERENCES.-Skeleton: Haddon 1879.

Subgenus tcymatholous Clark

Cgmatholcus Clark 1932, p. 132.

GENOTYPE.-Cymatholous longus Clark 1932.
TypE LOCALrrY AND HORIZON.-Hoot Owl Canyon, 15 mi. southwest

Vernal, Uinta County, Utah, U.S.A.; Duchesnean faunal age, Late Eocene.
DEFINITION.-An extinct North American subgenus of Geochelone

that derives its name from its characteristically strong looped sulcus be-
tween the marginal and plastral scutes; shell longer and higher in pro-
portion to width than other tortoises; lip conforming to contours of an-
terior lobe of plastron, bridge short, posterior lobe long.

GEOLOGIC RANGE.-Late Eocene.
GEOGRAPHIC RANGE.-Utah and Alabama, U.S.A.
REMARKS.-Certainly close to the subgenus Hadrianus of Geochelone,

but considerably more specialized. Placed as subgenus of Geochelone
by Auffenberg ( 1971).

Geochelone (Cymatholcus) flongus Clark
Cymatholcus Zongus Clark 1932, p. 132.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 153

TypE.-Carnegie Museum; a complete shell and parts of the skeleton.
TYPE LOCALITY AND HORIZON.-Hoot Owl Canyon, 15 mi. southwest

Vernal, Uinta County, Utah, U.S.A.; Duchesnean faunal age, Late Eo-
cene.

GEOLOGIC R~NCE.-Late Eocene.
GEOGRAPHIC RANGE.-Eastern Utah.

Geochelone ( Cymatholcus) tschucherti (Hay)

Hadrianus schucherti Hay 1902b, p. 22, pls. 4-5.
?Hadrianus schucherti Hay 1908, p. 382.
Cumatholcus schucherti Williams 195Oa, p. 30.

TYPE.-U. S. National Museum; a shell.
TypE LOCALITY AND HORIZON.-Near the Cocoa Post Office, Choc-

taw County, Alabama, U.S.A.; Duchesnean faunal age, Late Eocene.
GEOLOGIC RANGE.-Late Eocene.
GEOGRAPHIC RANGE.-Western Alabama, U.S.A.
REMARKS.-Williams ( 195Oa) placed schucherti in the genus Cyma-

tholcus on the basis of similarities in the details of form and the sulcal
relationships of the plastral elements. He suggested schucherti may
even be conspecific with longus.

Subgenus Geochelone Fitzinger

G€ochelone Fitzinger 1835, p. 108, 112, 122, ( as a subgenus) ( T. stellata Schweigger
[=T. elegans Schoepff] ).

Centrochelys Gray 1872a, p. 5 ( T. sulcata Miller).
Stigmochelys Gray 1873a, p. 5 (T. pardalis Bell).
Megachersine Hewitt 1931, p. 257 ( T. pardatis Bell ).

TYPE SPECIES.-Testudo elegans Schoepff C - Geochelone [Geoche-
lone] elegans [Schoepff] ).

DEFINITION.-An Old World subdivision of the genus Geochelone in
which the external nares are not higher than wide; nuchal scute absent
( except in platynota) ; first dorsal vertebra short; supracaudal scute un-
divided; gulars paired but not divergent; entoplastron not crossed by
humeropectoral sulcus.

GEOLOGIC RANGE.-Tertiary of Europe.
GEOGRAPHIC RANGE.-Africa, Ceylon, India, and Burma.
REMARKS.-The giant fossil tortoises of Malta, Menorca, and Tene-

rife probably belong to this subgenus. Though it is widely distributed
at present, it had an even greater range in the past. It contains four liv-
ing species, G. pardalis, G. sulcata, G. elegans, and G. platynota. Skel-
etal references to these forms are Siebenrock 1900, Deraniyagala 1930,
1939, Williams 195Ob, Auffenberg 1966b. Fifteen extinct species are
known.



154 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

Geochelone (Geochelon€) tamberiacensis ( Deperet)

Testudo amberiacends Deperet 1894, p. 717, pl. 24.

TypE.-The Faculty of Science ( Lyons); an entoplastron and nuchal
bone.

TYPE LOCALITY AND HORIZON.-Amb6rieu, Bresse, Ain Department,
France; Pontian faunal age, Early Pliocene.

GEOLOGIC RANGE.-Early Pliocene.
GEOGRAPHIC RANGE.-France.
REMARKS.-A poorly defined species without a nuchal scute. Ac-

cording to Szalai ( 1934) it is similar to Testudo kalksburgensis, but this ~
is probably in error as it seems to be a member of the genus Geochelone.

Geochebne (Geochelone) fammon ( Andrews)

Testudo ammon Andrews and Beadnell 1903, p. 5 (nomen nudum).
Testudo ammon Andrews 1904, p. 529.

TypE.-Geological Museum ( Cairo ); a complete shell.
TYPE LOCALITY AND HORIZON.-North of Birket-el-Qurun, Fayum,

United Arab Republic; Bartonian faunal age, Late Eocene.
GEOLOGIC RANGE,-Late Eocene.
GEOGRAPHIC RANGE.-Type locality.

REMARKS.-For best description, see Andrews ( 1906).

Geochelone ( Geochelone) tbeadnelli ( Andrews)

Testudo beadneli Andrews 1906, p. 285, fig. 91.

TYPE.-Geological Museum ( Cairo ); a complete shell.
TYPE LOCALMY AND HORIZON.-North of Birket-el-Qurun, Fayum,

United Arab Republic; Bartonian faunal age, Late Eocene.
GEOLOGIC RANGE.-Late Eocene.
GEOGRAPHIC R~NGE.-Type locality.
REMARKS.-Probably conspecific with G. ammon.

Geochelone (Geochelone) tbolivari ( Hernandez-Pacheco)

Testudo sp. Hernandez-Pacheco 1917, p. 197.
Testudo botivari Hernandez-Pacheco 1921, p. 329, 2 figs.
Testudo richardi Bergounioux 1938a, p. 62 ( part).

TypE.-Museo Nacional de Ciencias Naturales ( Madrid); an incom-
plete shell.

TYPE LOCALITY AND HORIZON.-Alcala de Henares, Palencia, Spain;

Miocene.
GEOLOGIC RANGE.-Miocene to ?Pliocene.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 155

GEOGRAPHIC RANGE.-Spain.
REMARKS.-It seems doubtful that aII the specimens Bgured by Royo

y Gomez ( 1935, fig. 5 ) as bolivari are the same species. Bergounioux
( 1938a) refers some to Testudo richardi. Peyer ( 1942) suggests that G.
bolivari may be subspecifically related to G. uitodurana Biedermann.
The best description is by Royo y Gomez ( 1935).

Geochelone (Geochelone) f burchardi (Ahl)

Testudo burchardi Ahl 1926, p. 575, fig. 1.
Testudo buchardi Szalai 1933, p. 156 ( typographical error) .

TypE.-Zoological Museum, Univ. of Berlin; a femur and humerus.
TypE LOCALITY AND HoRIzoN.-South part of Tenerife, Canary Is-

lands, Atlantic Ocean; Pleistocene volcanic tuff.
GEOLOGIC RANGE.-Pleistocene.
GEOGRAPHIC RANGE.-Type locality.

Geochelone (Geochelone) f crassa ( Andrews)
Testudo crassa Andrews 1914, p. 181.
Testudo crassa Szalai 1938, p . 162.

TypE.-British Museum ( Natural History); pieces ofthe shell.
TypE LoCALITY AND HoRIzoN.-Bed 31, Kachuku near Karungu,

Kenya, Africa; Burdigalian faunal age, Early Miocene.
GEOLOGIC RANGE.-Early Miocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-A very large tortoise unquestionably belonging in the

genus Geochelone.

Geochelone (Geochelone) tgrandis ( Macarovici and Vancea)

Testudo grandis Macarovici· and Vancea 1960, p. 381.

TypE.-Private collection of N. Macarovici, Malusteni, Romania;
fragments of the carapace and plastron.

TypE LOCALITY AND HORIZON.-Malusteni, Moldavia, Romania; As-
tian faunal zone, Late Pliocene.

GEOLOGIC RANGE.-Late Pliocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS .-Believed closely related to Testudo syrniiensis by Simi-

onesco ( 1930), but this is certainly incorrect.

Geochelone ( ?Geochelone) tgymnesica (Bate)
Testudo gymnesicus Hate 1914, p. 102, Bgs. 1-2.



156 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

TypE.-British Museum ( Natural History); portions of limbs.
TypE LoCALITY AND HORIZON.-Minorca Island; Late (?) Pleistocene.
GEOLOGIC RANGE.-Pleistocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-Probably related to the Pleistocene forms of Malta and

Africa, and here included within the subgenus Geochelone on the basis
of that supposition.

Geochelone (Geochelone) tisis ( Andrews)
Testudo isis Andrews 1906, p. 286, fig. 72.

TypE.-Geological Museum ( Cairo); an imperfect shell.
TypE LoCALITY AND HORIZON.-North of Birket-el-Qurun, Fayum,

United Arab Republic; Bartonian faunal age, Late Eocene.
GEOLOGIC RANGE.-Late Eocene.
GEoGRAPHIC RANGE.-Type locality.
REMARKS.-Relationships not clear.

Geochelone ( Geochelone) tmeshetica (Gabunya and Chkikvadze)
Testudo meschetica Gabunya and Chkikvadze 1960, p. 189, figs. 1-4.
Geochelone meschetica Chkikvadze 1970a, p. 59.

TYPE.-Geological collections, Academy of Sciences ( Georgia SSR);
partial shell.

TypE LOCALITY AND HORIZON.-Near Benar, Adigenski District,
Georgia, SSR; Middle or Late Oligocene.

GEOLOGIC RANGE.-Middle or Late Oligocene.
GEOGRAPHIC RANGE.-Georgia SSR.
REMARKS.-Close to G. ammon, according to Gabunya and Chkik-

vadze (1960).

Geochelone ( ?Geochelone) tnamaquensis ( Stromer)
Testudo namaquensis Stromer in Kaiser 1926, p. 139.

TypE.-Berlin Museum of Zoology; a complete plastron.
TypE LOCALrry AND HORIZON.-Namib, Namaqualand, South-west

Africa; Burdigalian faunal age, Middle Miocene.
GEOLOGIC RANGE.-Middle Miocene.
GEOGRAPHIC RANGE.-Type locality.

REMARKS.-Probably close to the extant Geochelone pardalis.

Geochelone ( Geochelone) pardalis (Bell)
Testudo pardalis Bell 1828, p. 420.
Testudo biguttata Cuvier 1836, p. 10 ( nomen nudum).



1974 AUFFENBERC: FOSSIL TORTOISE CHECKLIST 157

Teatudo bipunctata Gray 183la, p. 12 ( err6neously attributed to Cuvier MS).
Testudo armata Boil ( MS) (listed as synonym by Gray 193la, p. 4 ( nomen nudum).
Testudo pardalis Gray 183la, p. 12.
Geochelone pardatis Fitzinger 1835, p. 211.
Testudo sulcata Smets 1885, p. 8.
Homopus signatus Boettger 1893, p. 8 ( part).
Testudo calcarata Vaillant 1904, p. 186 ( part).
Megachersine pardalis Hewitt 1933, p. 257.
Testudo pardatis babcocki Loveridge 1935, p. 4.
Testudo pardalis pardalis Loveridge 1935, p. 4.
Geochelone pardalis Hewitt 1937, p. 789.
Testudo pardalis pardahs Mertens 1937, p. 5.
Geochelone pardalis babcocki Loveridge and Williams 1957, p. 235.
Geochelone paiudalia pardalis Loveridge and Williams 1957, p. 251.

TypE.-British Museum ( Natural History) ?; a preserved adult.
TYPE LOCALrry.-Mt. Debasien, Karamojo, Uganda, Africa.
GEOLOGIC RANGE.-The species is reported from Early Pleistocene

( Lehmann 1957) and Middle Pleistocene ( Broadley 1962) to Recent of
Africa.

GEOGRAPHIC RANGE.-G. p. babcocki ranges from Sudan and Ethopia
south to Natal, west through Cape of Good Hope Province to South-west
Africa, where it meets the typical form, Geochelone pardalis pardalis.

REMARKS.-For the most recent monographic treatment see Love-
ridge and Williams ( 1957). Two extant subspecies are recognized.

Geochelone ( Geochelone) perpiniana tleberonensis C Deperet)
Testudo teberonensis Deperet 1890, p. 915.
Testudo luberonensis Joleaud 1906, p. 360 ( typographical error).
Testudo perpiniana var. leberonensis Bergounioux 1936a, p. 28.

TypE.-Museum of Natural History ( Paris); part of carapace.
TYPE LOCALITY AND HoRIZ0N.-Mount Leberon, west of Cucuron,

France; Pontian faunal age, Early Pliocene.
GEOLOGIC RANGE.-Early Pliocene only? ( see remarks below).
GEOGRAPHIC RANGE.-Type 10Cality.
REMARKS.-Peyer ( 1942) considered leberonensis very close to, and

perhaps a synonym of Geochelone perpiniana, here treated as a separate
subspecies.

Geochelone ( Ceochelone) perpiniana tperpiniana ( Deperet)
Testudo perpiniana Deperet 1885, p. 214, pl. 4.

TypE.-Museum of Perpignon ( Southern France); slightly damaged
specimen.

TypE LoCALrrY AND HORIZON.-Near Perpignon, France; Plaisancian
faunal age, Late Pliocene.



158 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

~EOLOGIC RANGE.-Late Pliocene only? ( see remarks below).
GEOGRAPHIC RANGE.-France only? (see remarks below).
REMARKS.-G. perpiniana, G. picteti, G. bolivari, and G. uitodurana

all seem very close to the pardalis-sulcata group, and for this reason they
are all placed in the same subgenus. For the best description of G. per-
piniana see Deperet and Donnezan ( 1887, 1893-5). Lydekker ( 1889b)
placed G. perpiniana close to Geochelone atlas. Arambourg and Piveteau
( 1929) described a skull referred to this subspecies.

Geochelone ( Geochelone ) tpicteti ( Biedermann)
Testudo picteti Biedermann 1863, p. 18, pls. 2, 2a.

TypE.-Winterthur Museum ( Switzerland); an almost complete shell.
TYPE LOCALrrY AND HORIZON.-Near Winterthur, Switzerland; Upper

fresh water marls of Veltheim, Vindobonian faunal age, Late Miocene.
GEOLOGIC RANGE.-Late Miocene only? ( see remarks below).
GEOGRAPHIC RANGE.-Switzerland only?
REMARKS.-Roger ( 1902) suggested this species may be a synonym

of G. vitodurana.

Geochelone ( ?Geochelone) tpyrenaica ( Deperet and Donnezan)

Testudo purenaica Deperet 1885, p. 216 ( preliminary notice).
Testudo pyrenaica Deperet and Donnezan 1893-5, p. 155, pl. 16.

TYPE.-Museum of Natural History ( Paris ); a shell.
TYPE LOCALITY AND HORIZON.-Serrat Vaquar, Roussillon, France;

Astian Faunal age, Late Pliocene.
GEOLOGIC RANGE.-Pliocene.
GEOGRAPHIC RANGE.-France.

REMARKS.-Placed in the genus Geochelone, subgenus Geochelone,
on the basis of the absence of a nuchal scute. Szalai ( 1934) considered
it close to Testudo amiatae and Testudo marmorum ( of Gaudry), but
this is obviously incorrect.

Geochelone (?Geochelone) trichardi ( Bergounioux)

Testudo richardi Bergounioux 1938a, p. 271.

TYPE.-Geologic Museum, Seminario Conciliar de Barcelone; a
poorly preserved shell.

TYPE LOCALITY AND HORIZON.-Terrega, Lerida Province, Catalonia
Region, Spain; Oligocene.

GEOLOGIC RANGE.-Oligocene.
GEOGRAPHIC RANGE.-Catalonia Region, Spain.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 159

REMARKS.-Bergounioux ( 1938a) believed this species to be close to
Testudo castrensis.

Geochelone ( ?Geochelone) trobusta ( Leith-Adams)

Testudo fobusta Leith-Adams 1877, p. 178, pls. 5-6 ( not fobustus of Gilmore ).
Testudo sp,atti Leith-Adadis 1877, p. 186 ( Maghlak, Malta).
Testudo robustissima Tagliaferro 1914, p. 77 ( Corradino, Malta).

TypE.-British Museum ( Natural History); two vertebrae and parts
of the appendicular skeleton, including a tibia.

TYPE LOCALITY AND HORIZON.-Zebbug Cave, Malta; Middle Pleisto-
cene.

GEOLOGIC RANGE.-Middle Pleistocene.
GEOGRAPHIC RANGE.-Ta Xolea, Corradino Hill, Maghlak, Char

Dalam, and Zebbug Caves, Malta.
REMARKS.-A poorly defined species. Geochelone robustissima and

G. spratti are placed here because they differ only in size. This actioh
had apparently been suggested by C. Gatto during the discussion follow-
ing the Oral presentation by Tagliaferro ( 1914:79). Bones referrable to
all three species are known to occur in the same caves and in the same
deposits within them.

Geochelone ( ?Geochelone) tturgae ( Kuznetzov)

Testudo turgae Kuznetzov 1958, p. 66, figs. 1-2.
Testudo turmae Kuhn 1964, p. 32 ( typographical error).

TypE.-Location unknown to author; plastral fragments.
TypE LOCALITY AND HORIZON.-Turgai depression, Kazakhstan,

USSR; Miocene.
GEOLOGIC RANGE.-Miocene.
GEOGRAPHIC RANGE.-Kazakhstan, USSR.
REMARKS.-Here placed in the genus and subgenus Geochelone

solely on the basis of its rather large size. Additional material is needed
to determine its status and relationships.

Geochelone ( Geochelone) toitodurana ( Biedermann)

Testudo uitodurana Biedermann 1863, p. 13, pls. 1, la, 3.

TypE.-Winterthur Museum ( Switzerland); an almost complete shell.
TYPE LOCALITY AND HORIZON.-Near Winterthur, Switzerland; Up-

per fresh water bed of Veltheim, Vindobonian faunal age, Late Miocene.
GEOLOGIC RANGE.-Late Miocene.
GEOGRAPHIC RANGE.-Switzerland and France.
REMARKS.-For best description see Peyer ( 1942), who also sug-



160 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

gested that G. vitodurana may be only subspecifically distinct from G.
bolivari.

Subgenus 9 Hesperotestudo Williams
Hesperotestudo Williams 195Oa, p. 25 ( as subgenus ).

TYPE SPECIES.-Testudo osborniana Hay ( - Geochelone [Hespero-
testudo] osborniana [Hay] ).

DEFINmON.-An extinct Palearctic subdivision of the genus Geoche-
lone with narrow nuchal and pectoral scutes, limbs heavily armored, with
fused dermal ossicles forming a caudal buckler, supported by elongated
transverse processes of the caudal vertebrae.

GEOLOGIC RANGE.-( ?)Eocene to Pleistocene of North America and
'Tertiary" of Asia.

GEOGRAPHIC RANGE.-Nearctic and Palearctic.
REMARKS.-For most inclusive subgeneric description see Auffenberg

(1963).

Geochelone ( Hesperotestudo) talleni Auffenberg
Geochelone alteni Auffenberg 1966a, p. 877.

TypE.-Florida State Museum; shell of adult female.
TypE LOCALrrY AND HoRIZON.-MeGehee Farm, Newberry, Alachua

County, Florida, U.S.A.; Hemphillian faunal age, Middle Pliocene.
GEOLOGIC RANGE.-Middle Pliocene.
GEOGRAPHIC RANGE.-Several localities in central Florida, U.S.A.
REMARKS.-Auffenberg ( 1966a) considered it close to, and perhaps

ancestral to G. incisa.

Geochelone ( Hesperotestudo) farenivaga Hay
Testudo arenivaga Hay 1907, p. 16, figs. 6-8.
Geochelone arenicaga Auffenberg 1963, p. 93.

TYPE.-Carnegie Museum; pygal and one peripheral.
TypE LOCALITY AND HoRrzoN,-2 mi. N, Agate Springs, Sioux Coun-

ty, Nebraska, U.S.A.; lower Harrison beds, Arikareen faunal age, Early
Miocene.

GEOLOGIC RANGE.-Early Miocene.
GEOGRAPHIC RANGE.-WeStern Nebraska, U.S.A.
REMARKS.-Loomis ( 1909 ) described a skull, plastron, and limb

elements.

Geochelone (Hesperotestudo) fcampester (Hay)

Testudo campester Hay 1903, p. 627 ( nomen nudum).



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 161

Testudo campester Hay 1908, p. 455, fgs. 610-613.
Copherus campester Williams 1950a, p. 30.
Testudo rexroadensis Oelrich 1952, p. 301 ( Rexroad fm., Kansas).
Geochelone ferroadensis Auffenberg 1963, p. 93.

TypE.-American Museum of Natural History; a nearly complete
plastron and parts of the carapace.

TYPE LOCALITY AND HORIZON.-Near Mt. Blanco, Crosby County,
Texas, U.S.A.; Lower Blanco beds, Early Blancan faunal age, Late Plio-
cene.

GEOLOGIC RANGE.-Late Pliocene to early Pleistocene ( see Holman
1969).

GEOGRAPHIC RANGE.-Northwestern Texas to southwestern Kansas,
U.S.A.

REMARKS.-Auffenberg ( MS ) considers Geochelone ( Hesperotes-
tudo) rexroadensis a synonym of campester and closely related to G.
orthopygia.

Geochelone (Hesperotestudo) tequicomes (Hay)
Testudo equicomes Hay 1917b, p. 41, pls. 1 and 3.
Geochelone equicomes Auffenberg 1962c, p. 630.

TYPE.-U, S. National Museum; an epiplastron.
TypE LOCALITY AND HORIZON.-Cragin Quarry, Meade County, Kan-

sas, U.S.A.; Kingsdown Formati6n, Rancholabrean faunal age, Sangamon
Interglacial, Pleistocene.

GEOLOGIC RANGE.-( Sangamon) Late Pleistocene.
GEOGRAPHIC RANGE.-Kansas, U.S.A.
REMARKS.-(~OSe to Geochelone ( Hesperotestudo) turgida (see

Auffenberg 1962c, 1963).

Geochelone ( ?Hesperotestudo ) texornata C Lambe)
Testudo exomata Lambe 1906, p. 187, pl. 3, figs. 1-3.
Geochelone exomata Auffenberg 1962c, p. 635.

TypE.-Geological Survey of Canada; fragmentary pleural bones.
TYPE LOCALrrY AND HORIZON.-Bone Coulee, Cypress Hills, Assini-

boia, Saskatchewan Province, Canada; Chadronian faunal age, Early
Oligocene.

GEOLOGIC RANGE.-Early Oligocene.
GEOGRAPHIC RANGE.-Saskatchewan, Canada.
REMARKS.-The thick and strongly sculptured nature of the type

material suggests that the species is close to the Geochelone turgida line.
Very poorly defined.



162 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

Geochelone (Hesperotestudo) tfarri (Hay)

Testudo farri Hay 1908, p. 318, pl, 69.
Geochelone farri Auffenberg 1963, p. 94.

TYPE.-Princeton University; a shell.
TypE LOCALITY AND HORIZON.-Smith River Valley, Montana, U.S.A.;

Deep River Formation, Barstovian faunal age, Late Miocene.
GEOLOGIC RANGE.-Late Miocene.

GEOGRAPHIC RANGE.-Montana, U.S.A.
REMARKS.-Close to G. tedwhitei according to Williams .( 1953b).

Geochelone ( Hesperotestudo) tgilberti ( Hay)

Xerobates undata Gilbert 1898, p. 143, figs. 1-4.
Testudo gilberti Hay 1899b, p. 349. . .
Geochelone gitberti Auffenberg 1963, p. 94.

TypE.-Univ. of Kansas; a complete skull and lower jaw.
TYPE LOCALITY AND HORIZON.-Phillips County, Kansas, U.S.A.; Loup

Fork, Barstovian faunal age, Late Miocene.
GEOLOGIC RANGE.-Late Miocene.
GEOGRAPHIC RANGE.-Western Kansas, U.S.A.

REMARKS.-Probably a synonym of Geochelone osborniana. In the
G. thompsoni-G. angusticeps-G. osborniana-G. orthopygia evolution-
ary line according to Matthew ( 1924).

Geochelone ( Hesperotestudo) timpensa ( Hay)

Testudo impensa Hay 1908, p. 431, pls. 76, 77.
Testudo immensa Riabinin 1915, p . 10 ( typographical error ).
Geochelone impensa Auffenberg 1963, p. 88.

TypE.-American Museum of Natural History; a shell and most of
skeleton.

TYPE LOCALITY AND HORIZON.-Near the mouth of the Madison

River, Broadwater County, Montana, U.S.A.; Madison Valley Formation,
Barstovian faunal age, Late Miocene.

GEOLOGIC RANGE.-Late Miocene.
GEOGRAPHIC RANGE.-Montana, U. S.A.
REMARKS.-May be a synonym of Geochelone osborniana.

Geochelone (Hesperotestudo) tincisa (Hay)

Testudo incisa Hay 1916a, p. 46, pl. 3.
Gopherus incisa Williams 1950a, p. 30.
Ceochelone C Hesperotestudo ) incisa Auffenberg 1963, p. 82.

TYPE.-U. S. National Museum; a xiphiplastron.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 163

TypE LOCALrrY AND HORIZON.-Ocala Lime Company Quarry, near
Ocala, Marion County, Florida, U.S.A.; (?) Sangamon Interglacial, Pleis-
tocene.

GEOLOGIC RANGE.-Pleistocene.
GEOGRAPHIC RANGE.-Florida, USA.
REMARKS.-In Geochelone ( Hesperotestudo) turgida evolutionary

line ( Auffenberg 1962c, 1963).

Geochelone ( Hesperotestudo) tinusitata (Hay)
Testudo inusitata Hay 1907, p. 18.
Testudo innistata Szalai 1930, p. 354 ( typographical error).
Gopherus inusitata Williams 195Oa, p. 30.
Geochelone inusitata Auffenberg 1964a, p, 6

TYPE.-Carnegie Museum; left side of shell.
TYPE LOCALITY AND HORIZON.-Near Canyon Ferry, Broadwater

County, Montana, U.S.A.; Deep River Formation, Barstovian faunal age,
Late Miocene.

GEOLOGIC R~NGE.-Late Miocene.
GEOGRAPHIC RANGE.-Montana, U.S.A.
REMARKS.-Has only hexagonal neurals, but with a prominent epi-

plastral lip. Probably a synonym of G. osborniana.

Geochelone ( Hesperotestudo) tiohnstoni Auffenberg
Testudo Tugosa Johnston p . 47 ( manuscript name )
Geochelone iohnstoni Auffenberg 1962c, p. 627.

TypE.-Panhandle Plains Historic Museum ( Canyon, Texas ); a shell.
TYPE LOCALITY AND HORIZON.-Bed 4, Cita Canyon, Tule County,

Texas, U.S.A.; Late Blancan faunal age, Early Pleistocene.
GEOLOGIC RANGE.-Early Pleistocene.
GEOGRAPHIC RANGE.-Western Texas, U.S.A.
REMARKS.-In the Geochelone turgida evolutionary line ( Auffenberg

1962c, 1963).

Geochelone ( Hesperotestudo) tkalganensis ( Gilmore)
Testu(to kalganensis Gilmore 1931, p. 247, pl. 9.
Copherus kalganensis Williams 195Oa, p. 22-23.
Geochetone kalganensts Auffenberg 1962a, p. 633.

TypE.-American Museum of Natural History; partial shell.
TYPE LOCALITY AND HORIZON.-Kalgan area, Changchiaklou Hopeh

Province, China; Tertiary.
GEOLOGIC RANGE.-Tertiary, probably Pliocene or Pleistocene.
GEOGRAPHIC RANGE.-North China.



164 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

REMARKS.-Probably in the Geochelone turgida evolutionary line
( Auffenberg 1962c, 1963).

Geochelone ( Hesperotestudo) f klettiana ( Cope)
Testudo klettiana Cope 1875, p. 75.
Geochelone klettiana Auffenberg 1963, p. 94.

TYPE.-U. S. National Museum; a pygal.
TypE LOCALITY AND HoRIZON.-North of Santa Fe, New Mexico,

U.S.A.; "Loup Fork" Santa Fe series, Barstovian faunal age, Late Mio-
cene.

GEOLOGIC RANGE.-Late Miocene.
GEoGRAPHIC RANGE.-Type locality.
REMARKS.-Probably a synonym of Geochelone osborniana.

Geochelone ( Hesperotestudo) tniobrarensis ( Leidy)
Testudo ( Stylemys) niobrarensis Leidy 1858, p. 29.
Stulemys niobrarensts Cope 1870, p. 124.
Testudo niobrarensis Leidy 1873, p. 340.

TYPE.-U. S. National Museum; epiplastron and part of entoplastron.
TYPE LOCALITY AND HoRIZON.-"Niobrara River," Nebraska, U.S.A.;

( = Minnechaduza fauna?, Green 1956), Clarendonian faunal age, Early
Pliocene.

GEOLOGIC R~NGE.-PliOCene.
GEOGRAPHIC RANGE.-Southern South Dakota and northern Nebraska,

U.S.A.
REMARKS.-For best description, based on a more complete specimen

from Wolf Creek Fauna, Ogallala Formation, Late Clarendonian faunal
age, see Green ( 1956). Earlier references of this species to the Pleisto-
cene are in error.

Geoechelone ( Hesperotestudo) orthopygia tangusticeps ( Matthew)

Testudo angusticeps Matthew 1924, p. 207 ( in error).
Testudo orthopvgia angusticeps Matthew 1924, p. 210.
Geochelone angusticeps Auffenberg 1963, p. 89.

TypE.-American Museum of Natural History; complete skull and
greater part of shell, doubtfully associated.

TypE LOCALITY AND HORIZON.-Sheep Creek Quarry, Stonehouse
Draw, Sioux County, Nebraska, U.S.A.; Lower Sheep Creek beds, Late
Hemingfordian faunal age, Middle Miocene.

GEOLOGIC RANGE.-Middle Miocene.
GEOGRAPHIC RANGE.-Nebraska, U. S. A.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 165

REMARKS.-Perhaps close to the Geochelone osborniana evolution-
ary line. Matthew ( 1924) places it in the G. thompsoni-G. obsorniana
-G. gilberti-G. orthopygia line.

Geochelone ( Hesperotestudo) orthopygia forthopygia (Cope)
Xerobates orthopygius Cope 1878, p. 393.
Xerobates c!/clopygius Cope 1878, p. 394 ( "Miocene," Loup Fork, Kansas).
Caryoderma snouiana Cope 1866, p. 1044 ("Miocene," Loup Fork, Kansas).
Testudo undata? Williston 1898, p. 132.
Testudo orthopygia Hay 1899b, p. 349.
Testudo cyclopygia Hay 1899b, p. 349.
Testudo snouiana Hay 1902a, p. 451.
Geochelone orthopygia Williams 195Oa, p. 30.

TypE.-American Museum of Natural History; skull, jaw, plastron,
parts of carapace, and limb elements.

TYPE LOCALITY AND HORIZON.-Decatur County, Kansas, U.S.A.;
Republican River Formation, Clarendonian faunal age, Early Pliocene.

GEOLOGIC RANGE.-Pliocene.
GEOGRAPHIC RANGE.-Western Kansas and eastern Colorado, U.S.A.
REMARKS.-(lOSe to Geochelone ( Hesperotestudo) campester. In

the G. thompsoni-G. angusticeps-G. osborniana-G. gilberti line, ac-
cording to Matthew ( 1924). For best description of skeletons see Hay
(1908).

Geochelone (Hesperotestudo) tosborniana (Hay)
Testudo osbomiana Hay 1904b, p. 503.
Geochetone osborniana Williams 1950a, p. 30.

TypE.-American Museum of Natural History; a complete shell,
skull, and post cranial skeleton.

TYPE LOCALrry AND HoRIzoN.-Pawnee Creek, north of Sterling,
Weld County, Colorado, U.S.A.; Pawnee Creek Formation, Barstovian
faunal age, Late Miocene.

GEOLOGIC RANGE.-Late Miocene.
GEOGRAPHIC RANGE.-Northeastern Colorado, U.S.A.
REMARKS.-Probably includes Geochelone impensa and Geochelone

klettiana. In G. thompsons-G. angusticeps-G. orthopygia--G. gilberti
evolutionary line ( Matthew 1924).

Geochelone (Hesperotestudo) fprimaeua (Oelrich)
Testudo primaeua Oelrich 1950, p. 44.

TypE.-Univ. of Michigan Museum of Paleontology; shell and a few
girdle and limb elements.



166 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

TYPE LoCALrrY AND HORIZON.-Locality 8, west side Sweetwater
Creek, about 2 miles southwest Belmont Park Ranch House, N.E. 1/4
Sec. 32, TSS, R5W, Madison County, Montana, U.S.A.; Early Miocene.

GEOLOGIC RANGE.-Early Miocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-Relationship not clear.

Geochelone ( Hesperotestudo) triggai ( Hibbard)

Testudo riggsi Hibbard 1944, p. 72, fig. 1.
Gopherus riggsi Hibbard and Riggs 1949, p. 834.
Testudo turgida Oelrich 1957, p. 928 (part )
Geochelone riggsi Auffenberg 1962c, p. 628.

TypE.-Univ. of Kansas; a shell.
TypE LOCALI'rY AND HORIZON.-Seward County, Kansas, U.S.A.;

Rexroad Formation, Saw Rock Canyon fauna, Early Blancan faunal age,
Late Pliocene.

GEOLOGIC RANGE.-Late Pliocene.
GEOGRAPHIC RANGE.-Southwestern Kansas, and western Oklahoma,

U.S.A.
REMARKS.-In Geochelone turgida evolutionary line ( Auffenberg

1962c, 1963).

Geochelone ( Hesperotestudo) f turgida ( CoDe)

Emys turgida Cope 1876, p. 125.
Testudo turgida Cope 1892a, p. 127.
Gophems turgida Williams 1950a, p. 30.
Geochelone tufgida Auffenberg 1962c, p. 628.

TypE.-Academy of Natural Sciences ( Philadelphia ); a partial shell.
TypE LOCALITY AND HORIZON.-Three miles N. Dockum, near Es-

quella, Dickens County, Texas, U.S.A.; Ogallala Formation, Hemphillian
faunal age, Middle Pliocene ("Blanco Beds" in error by Cope 1892b).

GEOLOGIC RANGE.-Middle Pliocene.
GEOGRAPHIC RANGE.-Oklahoma, Kansas, and Texas, U.S.A.
REMARKS.-An early member of the G. turgida evolutionary line

( Auffenberg 1962c). For a good description and discussion of this spe-
cies see Oelrich ( 1957). Wilson ( 1950) stated that the type may be at
the University of Texas, though Oelrich ( 1957) located it in Philadelphia.

Geochelone ( Hesperotestudo) tundata ( Cope)

Testudo undata Cope 1875, p. 995 (p. 74 in sep.).
Geochelone undata Auffenberg 1963, p. 94.

TYPE.-U. S. National Museum; pygal and seven peripherals.



1974  AUFTENBERG: FOSSIL TORTOISE CHECKLIST 167

TypE LOCALrrY AND HORIZON.-Santa Fe Basin, near Santa Fe,
New Mexico, U.S.A.; Santa Fe series, probably Clarendonian faunal age,
Early Pliocene.
- GEOLOGIC RANGE.-Early Pliocene.

~EOGRAPHIC RANGE.-Central New Mexico, U.S.A.
REMARKS.-Cbsely related to Geochelone orthopygia.

Geochelone (Hesperotestudo) tuaga (Hay)
Testudo vaga Hay 1908, p. 414, pl. 19.
Gopherus vaga Williams 195Oa, p. 30.
Geochelone vaga Auffenberg 1964a, p. 6.

TypE.-American Museum of Natural History; apparently parts of
three individuals.

TYPE LOCALMY AND HORIZON.-Near Laramie Peak, Wyoming,
U.S.A.; Deep River Formation, Barstovian faunal age, Late Miocene.

GEOLOGIC RANGE.-Late Miocene.
GEOGRAPHIC RANGE.-Central Wyoming, U.S.A.
REMARKS .-Relationships not clear.

Geochelone ( Hesperotestudo) twilsoni ( Milstead)
Testudo wilsoni Milstead 1956, p. 168.
Geochelone wilsoni Aufferiberg 1962c, p. 630.

TYPE.-Texas Memorial Museum; a complete shell.
TYPE LOCALITY AND HORIZON.-Friesenhahn Cave, Bexar County,

Texas, U.S.A.; Rancholabrean faunal age, Wisconsin, Pleistocene.
GEOLOGIC RANGE.-Wisconsin Pleistocene td Early Recent.
GEOGRAPHIC RANGE.-Eastern Oklahoma and Central Texas to New

Mexico, U.S.A.
REMARKS.-Last known member of the imp6rtant Geochelone tur-

gida evolutionary line (Auffehberg 1962c, 1963).

Subgenus Indotestudo Lindholm
Testudo Schlegel and Miiller 1840, p. 30 ( part).
Indotestudo Lindholm 1929, p. 285 ( as subgenus).

TYPE SpECIES.-Testudo elongata Blyth ( = Geochelone [Indotestudo]
elongata [Blyth]).

DEFINmON.-An Asiatic subgenus of the genus Geochelone with a
low shell and no radiating pattern; nuchal may be either present or not,
entoplastron usually crossed by the humeropectoral sulcus.

GEOLOGIC RANGE.-Eocene to Recent.
GEOGRAPHIC RANGE.-Asia and the East Indies.



168 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

REMARKS.-A relatively primitive, tropical mesic forest group com-

prised of at least three extant and two extinct species. The extant popu-
lations are possibly all conspecific on the basis of broad overlap of char-
acters, as indicated by Smith ( 1931) and others. The living species are
elongata, travancorica, and forsteni.

REFERENCES.-Skeleton: Gray 1870c; Auffenberg 1966b.

Geochelone (?Indotestudo) tkaiseni (Gilmore)

Testudo kaiseni Gilmore 1931, p. 236, pl. 6.
Geochelone kaiseni Loveridge and Williams 1957, p. 224.
Geochelone kaisini Mlynarski 1968, p. 96 ( typographical error).

TypE.-American Museum of Natural History; a shell.
TypE L0CALrrY AND HORIZON.-Ardyn Obo Basin, Chinese Postroad,

Mongolian Republic; Ardyn Obo Formation, Late Eocene.
GEOLOGIC RANGE.-Late Eocene.
GEOGRAPHIC RANGE.-Mongolian Republic.
REMARKS.-May be close to, or included in Testudo antiqua group

according to Glaessner ( 1935), but this is not certain according to Mly-
narski (1955b). Mlynarski (1968) believes ithas noclose relatives.

Geochelone ( ?Indotestudo) tnana ( Gilmore)

Testudo nanus Gilmore 1931, p. 241, pl. 7.
Geochelone nana Loveridge and Williams 1957, p. 224.

TypE.-American Museum of Natural History; complete shell.
TYPE LOCALITY AND HoR]ZON.-East Mesa, Twin Oboes, Shara

Murun Region, Innef Mongoliap ?Ulan Gochu Formation, Early Oligo-
cene.

GEOLOGIC RANGE.-Early Oligocene.
GEOGRAPHIC RANGE.-Inner Mongolia.

REMARKS.-Several peculiar morphologic features of this species
suggest that its present subgeneric allocation may be incorrect.

Subgenus Manouria Gray

Testudo Schlegel and Muller 1840, p. 37 (part).
Geoemyda Cantor 1847, p. 2 ( part).
Manouria Gray 1852, 8 133.
Teleopus LeConte 1854, p. 187 ( Type T. luxatus).
Scapia Gray 1869, p. 169 (Type S. falconeri).
Emys Maack 1869, p. 278 (part).
Testudo Leidy 187la, p. 154 ( part).
Hadrianus Cope 1872a, p. 2.

TYPE SPECIES.-Testudo emys Schlegel and Muller ( -Geochelone
[Manouria] emys [Schlegel and Miiller]).



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 169

DEFINITION.-A rather primitive Holarctic and Oriental subgenus
of the genus Geochelone,,characterized by a long narrow skull with
many emyclid characters, a somewhat depressed shell with a nuchal scute,
divided supracaudal, narrowed pectorals ( wider in some fossil species ),
and a tendency to well developed flat epiplastral projections that are
often truncated and notched anteriorly on either side of the median line.

GEOLOGIC RANGE.-Eocene to Recent.
GEOGRAPHIC RANGE.-Extant species restricted to southeastern Asia.

Fossil distribution, Holarctic.
REMARKS.-A very primitive tropical mesic forest group, ancestral

to several subgenera and genera ( Auffenberg 1971). It contains two
extant ( G. impressa and G. emys) and seven extinct species. See addi-
tional comments under Kansuchelys. Hadrianus was originally consid-
ered a genus, then a subgenus of Testudo ( Williams 1953b), then of
Geochelone ( Loveridge and Williams 1957), then as a synonym of
Manouria ( Auffenberg 1971).

Geochelone ( Manouria) tcorsoni (Leidy)
Testudo corsoni Leidy 187la, p. 154.
Emys carteri Leidy 187lb, p. 372.
Testudo hadriana Cope 1872b, p. 5.
Testudo hadrianus Cope 1872d, p. 463.
Hadrianus octonaria Cope 1872b, p. 36.
Hadrianus corsoni Cope 1872b (1873), p. 36.
Hadrianus quadfatUs Cope 1872a, p. 468.
Hadrianus octonarius Cope 1872b, p. 3.

TypE.-Academy of Natural Sciences ( Philadelphia); the anterior
part of a plastron.

TypE LoCALITY AND HORIZON.-15 mi. S.E. Fort Bridger, Bridger
Basin, Wyoming, U.S.A.; Uinta Formation, Uintan faunal age, Late
Eocene.

GEOLOGIC RANGE.-Late Eocene.
GEOGRAPHIC RANGE.-Wyoming, U.S.A.

Geochelone ( Manouria) emys Schlegel and Muller)
Testudo emus Schlegel and Muller 1840, In Temminck, p. 34 ( Sumatra ) C see RE-

MARKS ~.
Geoemyda spinosa Cantor 1847, p. 2 ( part).
Testudo emydoides Dumeril and Dumeril 1851 ( substitute name for T. emys Schlegel

and Muller ).
Testudo phayrei Blyth 1863 ,p. 639.
Manouria fusca Gray 1852, p. 134 ( Singapore).
Teleopus luxatus LeConte 1854, p. 187 ( Java).
Manouria luxata Strauch 1862, p. 25.
Manouria emys Gunther 1864, p. 10.



170 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

Testudo ( Scapia) falconeri Gray 1869, p. 169 ( India?).
Scapia gigantea Gray 1872b, p. 8.
Manuria emys Lydekker 1889b, p. 209 ( Pliocene specimen) ( typographical error).

TypE.-Leiden Museum; a mounted adult.
TypE LocALrry.-Sumatra.
GEOLOGIC RANGE.-Pliocene of the Siwaliks ( Lydekker 1889b) to ~

the Recent.
GEOGRAPIDC RANGE.-Assam, Burma, Thailand, Malay Peninsula,

Sumatra, and Borneo.
REMARKS.-The publication dates of Temminck's works range from

1839-1844, but the number containing the description of T. emys Was
issued in 1840. The plastron of Geochelone emys reported from the
Shang dynasty ruins of Anyang, China ( Wu 1943) is undoubtedly the re-
sult of importation during occupation of the site. The Species is very
closely related to Geochelone impressa C Smith 1922, 1930). Lydekker
( 1889b) suggested Geoch€lone puniabiensis may be close to G. emys

REFERENCES.-Gray 1855, 1870c, Boulenger 1889, Jeude 1896, Wil-
liams 1950b, Auffenberg 1966b.

Geochelone ( Manouria) teocaenica ( Hummel)
Testudo eocaenica Hummel 1935, p. 463, pls. 2, 3.
Testudo eocenica Kuhn 1964, p. 117 C typographical error).

TypE.-Geological-Paleontological Museum, Univ. of Halle; a shell,
limbs, and skull.

TypE LoCALITY AND HORIZON.-Geisel Valley, Bavaria, Germany;
Auversian faunal age, Middle Eocene.

GEOLOGIC RANGE.-Middle Eocene.
GEOGRAPHIC RANGE.-Germany.
REMARKS.-The relationships are somewhat uncertain, but I agree

with Hummel ( 1935) and Zimmermann-Rollius ( 1966) that it is close
to G. emys.

Geochelone ( Manouria) tgilmorei ( new substitute name)
Hadrianus robustus Gilmore 1915, p. 146, pl. 25 ( preoccupied by Testudo robusta

Leith-Adams [1877], here placed in Geochelone [?Ceochelone] ).

TYPE.-Carnegie Museum; anterior half of plastron.
TypE LOCALITY AND HORIZON.-Near Kennedy's Hole, Uinta Basin,

Uinta County, Utah, U.S.A.; Horizon C, Uinta Formation, Uintan faunal
age, Late Eocene.

GEOLOGIC RANGE.-Late Eocene.
GEOGRAPHIC RANGE.-Utah, U.S.A.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 171

REMARKS.-Geochelone gilmorei is a new combination due to the
arrangement suggested here that H. robustus Gilmore and T. robusta
Leith-Adams belong to the genus Geochelone.

Geochelone ( ?Manouria) tinsolitus ( Matthew and Granger)
Testudo insolitus Matthew and Granger 1923, p. 5.
Testudo demissa Gilmore 1931, p. 232, figs. 11-15, pls. 5,8 ( Eocene, Outer Mon-

golia).
Testudo insolita Glaessner 1933, p. 282.
Geochelone insolitus Mlynarski 1968, p. 87, figs. 1-6.

TypE.-American Museum of Natural History; parts of carapace and
a left hypoplastron.

TYPE LOCALrrY AND HORIZON.-PfOmOntOry Bluff, Mongolian Repub-
lie; Ardyn Obo Formation, Late Eocene.

GEOLOGIC RANGE.-Late Eocene.
GEOGRAPHIC RANGE.-Mongolian Republic.
REMARKS.-Was thought to belong to the antiqua-graeca group

( Glaessner 1933, Mlynarski 1955b), but later redescribed and placed in
Geochelone by Mlynarski ( 1968). Here tentatively placed in the sub-
genus Manouria.

Geochelone ( ?Manouria) tmargae ( Hooijer)

?Testudo margae Hooijer 1948, p. 1169, p. 1.
Testudo margae Hooijer 1954, p. 486.

TypE.-Netherlands Museum of Natural History ( Leiden ); a right
scapula.

TYPE LOCALITY AND HoRIZON.-Desa Beru, Tjabenge ( Sopeng Dis-
trict) about 100 km. N.E. of Makasar, South Celebes, Indonesia; Pleisto-
cene.

GEOLOGIC RANGE.-Pleistocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-A large species of tortoise, presumed by Hooijer ( 1954)

t6 be related to some of the Upper Siwalik species, though this is un-
certain.

Geochelone ( Manouria) tmaiuscula (Hay)
(?) Hadrianus corsoni Cope 1874, p. 36 (part).
Hadrianus maiusculus Hay 1904a, p. 271, pl. 15.

TYPE.-Yale Peabody Museum; a shell.
TypE LOCALITY AND HORIZON.-Near Gallina, Rio Arriba County,

New Mexico, U.S.A.; Wasatchian faunal age, Early Eocene.
GEOLOGIC RANGE.-Early Eocene.



172 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

GEOGRAPHIC R~NGE.-New Mexico, U.S.A.
REMARKS.-The oldest known species of land tortoise. The ending

of the species name has been changed to correspond with the feminine
gender of Geochelone.

Geochelone ( Manouria) obailiensis Chkikvadze
Hadrianus obailinsis Chkikvadze 1970b, 751, Eg. 3 ( typographical error).

TypE.-Georgian Academy of Sciences ( Tbilisi); plastron.
TYPE LOCALITY AND HORIZON.-Georgian SSR, eastern Kazakhstan,

Zaisan Valley; Eocene, Obayla Formation.
GEOLOGIC RANGE.-Eocene.
GEOGRAPHIC RANGE.-Georgian SSR.
REMARKS.-Tentatively placed in the subgenus Manouria on the

basis of shell shape.

Geochelone ( ?Manouria) tpun~abiensis ( Lydekker)
Testudo puniabiensis Lydekker 1889b, p. 87.

TypE.-Indian Museum; right and left epiplastra.
TypE LoCALITY AND HoRIZON.-Siwalik Hills, Punjab, India; Plio-

cene (?).
GEOLOGIC RANGE.-Pliocene (P).
GEOGRAPHIC RANGE.-Punjab, India.
REMARKS.-Lydekker C 1889b) suggests it may be close to G. emys.

Geochelone ( Manouria) ttumida (Hay)
Hadrianus tumidus Hay 1908, 8 380.

TYPE.-American Museum of Natural History; a partial plastron and
peripherals.

TypE LOCALITY AND HoRIzoN.-Utah, U.S.A.; Uinta Formation, Uin-
tan faunal age, Late Eocene.

GEOLOGIC RANGE.-Late Eocene.
GEOGRAPHIC R~NGE.-Utah, U.S.A.
REMARKS.-The ending of the species name has been changed to

correspond with the feminine gender Geochelone.

Geochelone ( Manouria ) tutahensis C Gilmore)
Hadrianus utahensis Gilmore 1915, p. 148.

TYPE.--Carnegie Museum; plastron and bridge.
TypE LoCALITY AND HomzoN.-South of Kennedy's Hole, Uinta



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 173

County, Utah, U.S.A.; Horizon B or C, Uinta Formation, Uintan Faunal
age, Late Eocene.

GEOLOGIC RANGE.-Late Eocene.
GEOGRAPHIC RANGE.-Utah, U.S.A.

Subgenus f Megalochelys Falconer and Cautley
Megalochelys Falconer and Cautley 1837, p. 358.
Colossochelvs Falconer and Cautley 1844, p. 54.
Testudo Lydekker 1880, p. 20 (part).

TypE SpECIEs.-Colossochelys atlas Falconer and Cautley ( - Ceo-
chelone [ Megalochelys] atlas [Falconer and Cautley]).

DEFINITION.-An Old World subdivision of the genus Geochelone
in which the included species are large, thick-shelled tortoises with
swollen areas on the peripherals, nuchal scute absent, two gular scutes,
epiplastron strongly bifurcated anteriorly, and no narrowed pectorals.

GEOLOGIC RANGE.-Pleistocene and perhaps Recent.
GEOGRAPHIC RANGE.-Southeast Asia, East Indies, Mauritius Island,

Mascarene Group, Indian Ocean.
REMARKS.-An extinct, but specialized small group of large to giant

tortoises.

Geochelone (Megalochelys) tatlas ( Falconer and Cautley)
Megalochelys sivalensis Falconer and Cautley 1837, p. 358 ( subsequently withdrawn).
Colossochelvs atlas Falconer and Cautley 1844, p. 54.
Colossochel!/s (Megalochdys) atlas Maack 1869, p. 223.
Testudo ( Colossochelys) atlas Lydekker 1880, p. 20.
Testudo ( Megalochelys) atlas Lydekker 1889b, p. 209.
Testudo atlas Brown 1931, p. 183.
?Testudo ma,gae Hooijer 1948, p. 1169, fig. 1 ( Pleistocene, Celebes).
Testudo margae Hooijer 1954, p. 486.

TypE.-British Museum ( Natural History); an epiplastron.
TypE LOCALrry AND HoRIZON.-SiU'alik Hills, Punjab, India; Siwalik

Beds, Manchhar Series, Boulder-Conglomerate, Early Pleistocene.
GEOLOGIC RANGE.-Early to Late Pleistocene.
GEOGRAPHIC RANGE.-Punjab, India, Upper Burma, Java, Celebes,

and Timor.
REMARKS.-According to Lydekker ( 1889b) G. atlas is close to G.

pet'piniana. The 12-foot composite reconstruction of the type and as-
sociated specimens by Falconer is incorrect; it should be closer to 6 feet
( Lydekker 1889b). For latest discussion, see Hooijer ( 1971).

Geochelone (?Megalochelys ) 1 cautlegi (Lydekker)
Cautleva annuliger Theobald 1879, p. 186 (nomen nudum).



174 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

Testudo cautlet/i Lydekker 1889a, p. 86.
Testudo cauthley Riabinin 1915, p. 12 ( typographical error).

TYPE.-British Museum ( Natural History); an epiplastron.
TypE LoCALITY AND HoRIZON.-Siwalik Hills, near Nila, Potwar,

Punjab, India; Upper Siwalik beds, Potwar Silts, Middle Pleistocene?
GEOLOGIC RANGE.-Middle Pleistocene.
GEOGRAPHIC RANGE.-Punjab, India.
REMARKS.-Geochelone ( ?Megalochelys) cautleyi is smaller than G.

atlas and generally believed not the same species on the basis of the
available epiplastron. The differences are conceivably sexually corre-
lated.

Geochelone ( Megalochelys) tgadowi (Van Denburgh)

Testudo guentheri Gadow 1894, p. 320 ( invalid name, not of Baur 1889, proposed
for T. st:meirei; Sauzier [part] ).

Testudo gadowi Van Denburgh 1914, p. 257.

TypE.-Museum of Zoology, Cambridge University; anterior part of
plastron.

TYPE LOCALITY AND HORIZON.-Mare aux Songes, Mauritius Island,
Mascarene Group, Indian Ocean; Late Pleistocene or Early Recent.

GEOLOGIC RANGE.-Pleistocene and/or Recent.
GEOGRAPHIC RANGE.-Mauritius Island.

Subgenus tMonachelys Williams

Monachetvs Williams 1952, p. 547 ( as subgenus).

TYPE SPECIES.-Testudo ( Monachelus) monensis Williams ( = Geo-
chelone [Monachelys] monensis [Williams]).

DEFINITION.-A Nearctic subdivision of the genus Geochelone in
which the majority of the characters are as in the subgenus Chelonoidis,
except that the centrum of the Erst dorsal vertebra is very elongate and
the xiphiplastral notch is absent.

GEOLOGIC RANGE.-Known only from the Late (?) Pleistocene.
GEOGRAPHIC RANGE.-Mona Island, West Indies.
REMARKS.-The relationships of this monotypic subgenus are not

clear, but it is believed to be close to Chelonoidis.

Geochelone (Monachelys) tmonensis ( Williams)

Testudo ( Monachelys ) monensis Williams 1952, p. 547, pls. 44-47.

TypE.-American Museum of Natural History; skull, parts of shell,
and partial skeleton.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 175

TYPE LOCALITY AND HoRIZON.-Lirio Cave, Mona Island, West In-
dies; Late PIeistocene.

GEOLOGIC RANGE.-Late Pleistocene.
GEOGRAPHIC RANGE.-Mona Island, West Indies.

Subgenera Inquirendae

The following species belong to the genus Geochelone, but their
subgeneric placement is not clear.

Geochelone fbessarabica (Riabinin)

Testudo bessarabica Riabinin 1915, p . 6 , pls . 1 , 3, 4 .

TypE.-Geology Museum, Academy of Sciences ( Leningrad); partial
shell.

TypE LOCALrrY AND HORIZON.-Tarakliya, Bender District, Bessarabia
Province ( Cahul), Southern Moldavian S.S.R.; Meotian faunal age, Early
Pliocene.

GEOLOGIC RANGE.-Early Pliocene.
GEOGRAPHIC RANGE.-Central Bessarabia, Moldavian S.S.R., and

Moldavia, Romania.
REMARKS.-The best description of the shell is that by Macarovici

( 1930) from near the type locality. According to Riabinin ( 1915) it has
close affinities to Geochelone puniabiensis and G. emys. Glaessner ( 1933)
states that it is not in the antiqua-graeca phyletic line. Mlynarski ( 1969b)
correctly places it in Geochelone.

?Geochelone tchaileoti ( Bergounioux)

Testudo chaileoti Bergounioux 1935, p. 93, pl. 5.
Testudo chailloti Kuhn 1964, p. 114 (typographical error).

TypE.-Museum of Natural History ( Montauban, France ); a shell.
TypE LOCALrrY AND HORIZON.-Dieupentale, Tarn et Garonne Dept.,

France; Stampian faunal age, Middle Oligocene.
GEOLOGIC RANGE.-Middle Oligocene.
GEOGRAPHIC RANGE.-France.
REMARKS.-Presumably close to Testudo catalaunica and T. p!/renaica

according to Bergounioux ( 1935). The absence of a nuchal scute indi-
cates very clearly that this species belongs to Geochelone.

Geochelone f costaricensis ( Segura)

Testudo costaricensis Segura 1944, p. 9-29.
Geochelone costaricensis Loveridge and Williams 1957, p. 224.



176 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

TypE.-National Museum of Costa Rica; an almost complete shell.
TypE LOCALrrY Aive HoRIzoN.-Milla 52, Peralta District, Costa

Rica; Late Oligocene or Early Miocene.
GEOLOGIC RANGE.-Late Oligocene or Early Miocene.
GEOGRAPHIC RANGE.-COSta Rica.
REMARKS.-According to Segura ( 1944) it is close to Stylemys

amphithorax but Auffenberg ( 1971) suggests it is close to Indotestudo
and Chelonoidis.

?Geochelone f cultrata ( Cope)
Testudo cultratus Cope 1873a, p. 6.
Testudo cultrata Hay 1902a, p. 451.
Geochelone cultrata Auffenberg 1963, p. 94.

TypE.-U.S. National Museum; partial epiplastron.
TypE LoCALrrY AND HORIZON.-Head of Horse Tail Creek, Weld

County, Colorado, U.S.A.; White River Series, Chadronian fauhal age,
Early Oligocene.

GEOLOGIC RANGE.-Early Oligocene.
GEOGRAPHIC RANGE.-Northeastern Colorado, U.S.A.

?Geochelone f larteti Pictet
Testudo gigantea Lartet 1851, p. 38 ( preoccupied, not of Bravard).
Testudo larteti Pictet 1853, p. 444 ( substitute name for T. gigantea Lartet).
Testudo larteti Lydekker 1889a, p. 90.

TypE.-Location unknown to author; part of the shell.
TypE LOCALITY AND HoRIzoN.-Sansan, Gers Department, France;

Vindobonian faunal age, Late Miocene.
GEOLOGIC R~NGE.-Late Miocene.
GEOGRAPHIC RANGE.-Sansan, France.
REMARKS.-A giant tortoise, probably referable to the genus Geo-

chelone.

?Geochelone toskarkuhni Mlynarski
Ceochelone oskarkuhni Mlynarski 1968, p. 91, figs. 6-8, pl. 9.

TYPE.-Palaeozoological Institute, Polish Academy of Sciences ( War-
saw ); a plastron and posterior portion of carapace.

TypE LoCALITY AND HORIZON.-Alta Teli, Dzereg Valley, Western
Mongolia; Pliocene.

GEOLOGIC RANGE.-Pliocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-I agree with Mlynarski ( 1968) and others that the entire



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 177

question of relationships among the Asiatic Pliocene fossil tortoises needs
critical review. Until this is done the allocation of this species to Ceo-
chelone will remain in doubt.

?Geochelone tperagrans ( Hay)
Testudo peragrans Hay 1907, p. 15.
Geochelone peragrans Auffenberg 1963, p. 94.

TYPE.-Carnegie Museum; skull and shell.
TYPE LOCALITY AND HORIZON.-South of McCartney Mountain and

Big Hole River, north of Dillon, Montana, U.S.A.; (?) Oligocene.
GEOLOGIC RANGE.-( p)Oligocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-The relationships of this species are unknown.

?Geochelone tphosphoritarum ( Bergounioux)
Testudo phosphoritamm Bergounioux 1935, p. 85, pl. 14.

TYPE.-Toulouse Natural History Museum; partial carapace.
TypE LOCALITY AND HORIZON.-Quercy Region, France; Phosphorites

de Quercy, Oligocene.
GEOLOGIC RANGE.-Oligocene.
GEOGRAPHICAL RANGE.-Quercy Region, France.
REMARKS.-Close to Testudo gigas Bravard ( nomen nudum) ac-

cording to Bergounioux ( 1935).

?Geochelone fquadrata (Cope)
Testudo quadratus Cope 1885, p. 762, pl. 61.
Testudo quadrata Hay 1902a, p. 451.
Geochelone quadrata Auffenberg 1963, p. 94.

TypE.-American Museum of Natural History; and epiplastral pro-
jection.

TypE LOCALITY AND HORIzoN.-Head of Horse Tail Creek, Weld
County, Colorado, U.S.A.; White River Formation, Chadronian faunal
age, Early Oligocene.

GEOLOGIC RANGE.-Early Oligocene.
GEOGRAPHIC RANGE.-Southeastern Wyoming and northeastern Col-

orado, U.S.A.
REMARKS.-The relationships of this species are unknown.

?Geochelone tscha#eri ( Szalai)
Testudo schaiTeri Szalai 1933, p. 154 ( not Ptychogaster schalleri Glaessner ).



178 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

TYPE.-Vienna Museum of Natural History; a skull and femur.
TypE LOCALITY AND HORIZON.-Samos, Greece; Pontian faunal age,

Early Pliocene.
GEOLOGIC RANGE.-Early Pliocene.
GEOGRAPHIC RANGE.-Samos, Greece.

REMARKS.-A large tortoise, probably in the genus Geochelone,
subgenus Geochelone. Bachmayer ( 1967) described additional material
from Pikermi.

?Geochelone f sloanei ( Lydekker)

Testudo sloanei Lydekker 1889a, p. 89.

TypE.-British Museum ( Natural History ); a shell.
TypE LOCALITY AND HORIZON.-"Tertiary of Turkey."
GEOLOGIC RANGE.-Tertiary, probably early Pliocene.
GEOGRAPHIC RANGE.-Turkey.
REMARKS.-(10Sely related to Geochelone radiata or G. pardalls ac-

cording to Lydekker ( 1889a) though certain of its features suggest it
may be close to Testudo.

?Geochelone ttarakliensis Riabinin

Testudo tarakliensis Riabinin 1915, p. 12.
Testudo tarabliensis. Kuhn 1964, p. 134 ( typographical error).

TypE.-Geology Museum of the Academy of Sciences; carapace.
TYPE LOCALITY AND HORIZON.-Tarakliya, Bessarabia Province,

Moldavian S.S.R.; Meotian faunal age, Early Pliocene.
GEOLOGIC RANGE.-Early Pliocene.
GEOGRAPHIC RANGE.-Bessarabia, Moldavian S.S.R.
REMARKS.-AfEnities not clear. Not in the antiqua-graeca phyletic

line according to Glaessner ( 1933) and Mlynarski ( 1955a). Absence of
a nuchal scute suggests it belongs to the genus Geochelone.

Geochelone f thompsoni ( Hay)

Testudo thompsoni Hay 1908, p. 400, pl. 66.
Geochelone thompsoni Auffenberg 1963, p. 94.

TypE.-American Museum of Natural History; a skull and part of
plastron, cervical vertebrae and parts of foreleg.

TypE LOCALITY AND HORIZON.-Corral Draw, Ziebach County, South

Dakota, USA.; Lower Oreodon beds, Orellan faunal age, Middle Oligo-
cene.

GEOLOGIC RANGE.-Middle Oligocene.
GEOGRAPHIC RANGE.-Western South Dakota.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 179

REMARKS.-Though the exact relationships of this species remain
unknown, it is apparently in the phyletic line with Geochelone o. angus-
ticeps. G. osborniana, G. gilberti, and G. orthopygia, according to Mat-
thew ( 1924).

?Geochelone fulanensis ( Gilmore)
Testudo ulanensis Gilmore 1931, p. 245.
Geochetone ulanensis Mlynarski 1968, p. 96.
Hadfianus ulanensis Chkikvadze 197Ob, p. 71.

TypE.-American Museum of Natural History; part of plastron and
carapace.

, TYPE LOCALITY AND HORIZON.-North Mesa, Shara Murun Region,
Inner Mongolia, China; Ulan Shireh Formation, Ludian faunal age, Late
Eocene.

GEOLOGIC RANGE.-Late Eocene.
GEOGRAPHICAL RANGE.-Shara Murun Region, Inner Mongolia, China.
REMARKS.-AfAnities not clear but certainly not in the antiqua-

graeca phyletic line according to Glassner ( 1933 ). The material is very
fragmentary and detailed comparison is impossible. Chkikvadse ( 1907b)
places it in Hadrianus, but he does not state on what basis he does so.

?Geochelone tgunnanenals (Yeh)
Testudo yunnaneensis Yeh 1963a, p. 47, figs. 27-29, pls. XVI, XVII:
Testudo lunanensis Yeh 1963a, p. 50, pl. XIII ( Lower Oligocene, China).

TypE.-Institute of Paleontology and Paleoanthropology ( Peking);
fragmentary plastron.

TypE LOCALITY AND HORIZON.-Wa-yao-chung, Ta-i-ma, Lunan.
Yunnan, China; Lower Oligocene.

GEOLOGIC RANGE.-Lower Oligocene.
GEOGRAPHIC RANGE.-Type locality only.
REMARKS.-Testudo lunanensis Yeh is placed in the synonomy of T.

yunnanensis Yeh because of morphological similarity. Both are pro-
visionally placed in the genus Geochelone on the basis of their large size.

Genus Gopherus Raftnesque
Testudo Bartram 1791, p. 18.
Copherus Rafinesque 1832, p. 64.
Xerobates Agassiz 1857, p . 446.
Bysmachelys Johnston 1937, p. 439.

GENoTypE.-Testudo poll/phemus Daudin ( = Gopherus polyphemus
[Daudin]).

DE:FINmON.-Nearctic genus of tortoises with short cervical verte-



180 BULLETIN FL0RIDA STATE MUSEUM Vol. 18, No. 3

brae; median premaxillary ridge; flattened forelimbs adapted for digging;
nuchal scute usually as wide as long; hyoplastron usually longer than
hypoplastron; fourth vertebral scute usually wider than long.

GEOLOGIC RANGE.-Oligocene to Recent.
GEOGRAPHIC RANGE.-As a fossil almost all of the Nearctic region

south of Canada and south throughout northern Mexico to the State of
Aguascalientes. Recent range much smaller.

REMARKS.-For best generic description see Williams ( 195Oa). Rec-
ognition of this genus in the Tertiary of Asia by Williams ( 1952) was
based on (1) a presumed close relationship between turgida and kalgan-
ensis ( probably correct ), and (2) reference of turgida to the genus
Gopherus ( certainly incorrect). For further discussion see Oelrich
( 1957) and Auffenberg ( 1962c). Four extant species are recognized:
G. agassizii, G. berlandieri, G. Bavomarginatus, and G. polyphemus.

REFERENCES.-Zoogeography Blair 1958, Brattstrom 1961. Mor-
phology. Skeleton: Williams 195Ob, Auffenberg 1964b, 1966b, in press;
Skull: Williams 195Oa, Auffenberg in press.

Gopherus agassizii ( Cooper)

Xerobates agassizii Cooper 1863, p. 120.
Testudo agassizii Boulenger 1889, p. 156.
Gopherus agassizii Stejneger 1893, p. 161.
Gopherus polyphemus agassizii Mertens and Wermuth 1955, p. 351.

TypE.-California State Geological Survey; 3 young specimens.
TypE LoCALITY.-Near Fort Mojave, California, U.S.A.
GEOLOGIC RANGE.-Known from the Pleistocene of California ( Miller

1942, Brattstrom 1953, 1958), and the Post-Pleistocene of Nevada ( Bratt-
strom 1954).

GEOGRAPHIC RANGE.-Southwestern United States ( extreme south-
western Utah, southern Nevada, western Arizona, extreme northwestern
Baja California, southeastern California), and all of Sonora, Mexico to
extreme northern Sinaloa.

REMARKs.-Mertens ( 1956 ) records hybrids between this species
and G. berlandieri in captivity. Most closely related to G. berlandieri
( Auffenberg in press).

Gopherus tatascosae ( Hay)
Testudo atascosae Hay 1902c, p. 383.
Gopherus atascosae Hay 1924, p. 247.

TypE.-Academy of Natural Sciences ( Philadelphia); parts of a
plastron.



1974 AUFFENBERG: FOSSIL TORTOISE  CHECKLIST 181

TypE LocALrry AND HORIZON.-Atascosa County, Texas, U.S.A.;
probably Middle Pleistocene.

GEOLOGIC RANGE.-(?) Middle Pleistocene.
GEOGRAPHIC RANGE.-Central Texas, U.S.A.
REMARKS.-In 1908 Hay thought the type may have been taken

from Miocene beds, but he established its true origin in 1930. Closely
related to Gopherus polyphemus and perhaps conspecifc with it.

Gopherus fbrattstromi (new substitute name)
Gopherus depressus Brattstrom 1961, p. 548 ( Preoccupied by Testudo depressa

Guerin-Meneville [1829], here placed in Copherus polyphemus).

TypE.-California Institute of Technology; a nearly entire shell.
TYPE LOCALITY AND HORIZON.-Tehachapi Mountains, Kern County,

California, U.S.A.; Bopesta Formation, Cache Peak fauna, Barstovian
faunal age, Late Miocene.

GEOLOGIC RANGE.-Late Miocene.
GEOGRAPHIC RANGE.-Southeastern California, U.S.A.
REMARKS.-Close to G. mohavetus and G. pa·naa, according to Bratt-

strom ( 1961).

Gopherus fbrevisternus ( Loomis)
Testudo brevisterna Loomis 1909, p. 21.
Gopherus bret>istema Williams 1950a, p. 19.

TypE.-Amherst College; most of the entire skeleton.
TYPE LOCALITY AND HORIZON.-Muddy Creek, Laramie County,

Wyoming, U.S.A.; Upper Harrison beds, Arikareean faunal age, Early
Miocene.

GEOLOGIC RANGE.-Early Miocene.
GEOGRAPHIC RANGE.-Eastern Wyoming, U.S.A.
REMARKS.-The ending of the species name has been changed to

correspond with the masculine gender of Gopherus.

Gopherus f canyonensis C Johnston)
Bymnachelys cant/onensis Johnston 1937, p. 440.
Copherus can!/onensis Williams 195Oa, p. 21.

TypE.-Panhandle Historic Plains Museum ( Canyon, Texas ); a plas-
tron and skeleton ( plastron now lost).

TypE L0CALITY AND HoRIZON.-North Cita Canyon, Sec. 164, block
6, Randall County, Texas, U.S.A.; Cita Canyon beds, Late Blancan faunal
age, Early Pleistocene.

GEOLOGIC RANGE.-Early Pleistocene.



182 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

GEOGRAPHIC RANGE.-Northwestern Texas and beds of similar age
as far west as southern Arizona, U.S.A.

REMARKs.-Close to G. pertenuis. Close to G. polyphemus accord-
ing to Williams ( 1952).

Gopherus tcopei (Koerner)

Testudo copei Koerner 1940, p. 838.
Gophems copei Williams 195Oa, p. 30.

TYPE.-Yale Peabody Museum; a shell.
TypE LOCALITY AND HORIZON.-See. 14, RlON, TSE, Meagher Coun-

ty, Montana, U.S.A.; Deep River Formation, Barstovian faunal age, Late
Miocene.

GEOLOGIC RANGE.-Late Miocene.
GEOGRAPHIC RANGE.-Type locality.

REMARKS.-Close to G. emiliae, according to Koerner ( 1940), and
probably conspecific with it.

Gopherus fdehiscus Des Lauriers

Copherus dehiscus Des Lauriers 1965, p. 1.

TypE.-Los Angeles County Museum; internal cast of entire shell,
except anterior lip of carapace.

TypE LOCALITY AND HORIZON.-Cajon Pass, W. end of Cajon Valley,
NW-1/4 Sec. 1, N.E.-1/4 Sec. 2, T3N, R7W, SBB and M. San Antonio
Quad. 1/2 mile S.W. of Hwy. to Big Pines Recreation Area, San Berna-
dino County, California, U.S.A.; Barstovian faunal age, Late Miocene.

GEOLOGIC RANGE.-Late Miocene.
GEOGRAPHIC RANGE.-Type locality.

REMARKs.-Occurred sympatrically with the type of Geochelone
milleri ( Brattstrom).

Gopherus tedae (Hay)

Testudo edae Hay 1907, p. 19.
Gopherus edae Williams 195Oa, p . 30.

TYPE.-Carnegie Museum; most of a carapace and plastron.
TYPE LOCALITY AND HORIZON.-Near Running Water Creek, Sioux

County, Nebraska, U.S.A.; Harrison Formation, Arikareean faunal age,
Early Miocene.

GEOLOGIC RANGE.-Early Miocene.
GEOGRAPHIC RANGE.-Western Nebraska, U.S.A.
REMARKS.-Probably close to G. holtandi. The best description is by

Hay ( 1908).



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 183
Gopherus temiliae (Hay)

Testudo emiliae Hay 1908, p. 419.
Cophems emiliae Williams 1950a, p. 30.

TypE.-American Museum of Natural History; an almost entire shell.
TypE LoCALrry AND HORIZON.-On Porcupine Creek, South Dakota,

U.S.A.; Lower Rosebud Formation, Arikareean faunal age, Early Miocene.
GEOLOGIC RANGE.-Early Miocene.
GEOGRAPHIC RANGE.-Southern South Dakota, U.S.A.
REMARKS.-Presumably close to G. copei, according to Koerner

(1940).

Gopherus flauomarginatus Legler
Gopherus poluphemus Duges 1888, p. 146-147 ( part).
Copherus flauomarginata Legler 1959, p. 337.
Gophems polyphemus flauomarginatus Wermuth and Mertens 1961, p. 174.

TYPE.-U.S. National Museum; a mounted specimen.
TYPE LOCALITY.-30-40 miles north, of Ciudad Lerdo, Durango,

Mexico.
GEOLOGIC RANGE.-Middle Pleistocene and Recent.
GEOGRAPHIC RANGE.-In a living state known only from the Bolson

de Mapimi, northeastern Chihuahua, western Coahuila and northern
Durango, Mexico. Pleistocene fossils known from Aguascalientes, Mex-
ico.

REMARKS.-Most closely related to G. polyphemus C Legler 1959,
Auffenberg in press). Grant's statement ( 1960) that this species is
closest to G. agassizii is not followed here. Legler and Webb ( 1961)
described additional specimens.

REFERENCES.-Morphology. Skull: Legler 1959; Skeleton: Auffenberg
1966b, in press.

Gopherus thexagonatus (Cope)
Testudo hexagonata Cope 1893, p. 77.
Gopherus hexagonata Williams 195Oa, p. 30.

TypE.-Univ. of Texas; originally a partial shell, now represented
by a few badly broken pieces.

TYPE LOCALITY AND HORIZON.-Rock Creek, Tule Canyon, Briscoe
County, Texas, U.S.A.; Tule Formation, Rock Creek Local fauna, Irving-
tonian faunal age, Middle Pleistocene.

GEOLOGIC RANGE.-Middle to Late Pleistocene.
GEOGRAPHIC RANGE.-Most Of Texas, north to Kansas, U.S.A.
REMARKS.-For best  description see Auffenberg ( 1962b). The end-



184 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

ing of the species name has been changed to correspond with the mas-
culine gender of Gopherus. Probably a synonym of G. laticaudatus.

Gopherus thollandi (Hay)
Testudo hollandi Hay 1907, p. 18.
Gopherus hollandi Williams 1950a, p. 30.

TypE.-Carnegie Museum; a complete shell.
TypE LOCALrrY AND HORIZON.-Near Running Water Creek, Sioux

County, Nebraska, U.S.A.; Harrison Formation, Arikareean faunal age,
Early Miocene.

(EOLOGIC RANGE.-Early Miocene.
GEOGRAPHIC RANGE.-WeStern Nebraska, U.S.A.
REMARKS.-Probably close to G. edae.

Gopherus thuecoensis Strain
Copherus huecoensis Strain 1966, p. 24.

TypE.-Memorial Museum, Bureau of Economic Geology, Univ. of
Texas; plastron and various appendicular skeletal elements of same
individual.

TYPE LOCALrrY AND HORIZON.-Madden Arroyo, Hudspeth County,
Texas, U.S.A.; Fort Hancock Formation, Early Pleistocene.

GEOLOGIC RANGE.-Early Pleistocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-Clearly a member of the pol!/phemus group on the basis

of the carpus and probably a synonym of G. flavomarginatus

Gopherus flaticaudatus (Cope)
Testudo laticaudata Cope 1893, p. 75.
Testudo luticaudata Wilson 1950, p. 115 ( typographical error).
Geochelone Zaticaudata Auffenberg 1963, p. 93.

TypE.-Univ. of Texas; an epiplastron and xiphiplastron.
TYPE LOCALITY AND HORIZON.-Rock Creek, Tule Canyon, Briscoe

County, Texas, U.S.A.; Tule Formation, Rock Creek local fauna, Irving-
tonian age, Middle Pleistocene ( incorrectly stated as Pliocene by Hay
1908).

GEOLOGIC RANGE.-Middle Pleistocene.
GEOGRAPHIC RANGE.-Northwestern Texas, U.S.A.
REMARKS.-Gopherus laticaudatus is a new combination, based on

the similarity of laticaudatus and G. pertenuis. · The ending of the species
name has been changed to correspond with the masculine gender of
Gopherus. It is probably a synonym of G. hexagonatus.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 185

Gopherus flaticuneus (Cope)
Testudo laticunea Cope 1873a, p. 6.
Gopherus kticunea Williams 195Oa, p. 30.

TYPE.-American Museum of Natural History; an almost complete
carapace, pelvis, and some limb elements.

TypE LOCALI'rY AND HORIZON.-Head of Horsetail Creek, Weld
County, Colorado, U.S.A.; Horsetail Creek Member, White River Forma-
tion, Chadronian faunal age, Early Oligocene.

GEOLOGIC RANGE.-Early and Middle Oligocene.
GEOGRAPHIC RANGE.-Northeastern Colorado, southeastern Wyoming,

southwestern South Dakota, and western Nebraska, U.S.A.
REMARKS.-Copherus praeextans and G. neglectus may be synonyms

of G. latiouneus. Hay ( 1908 ) refers fossils from the Colorado Oreodon
beds to this species, but this stratigraphic assignment was based on an
error in geologic interpretation. The ending of the species name has
been changed to conform with the masculine gender of Copherus.

Gopherus tmohavetus ( Merriam)
Testu(to mohavense Merriam 1919, p. 456.
Testudo mohavensis Bataller 1926, p. 155.
Copherus mohavense Williams 195Oa, p. 30.
Copherus mohauetus Des Lauriers 1965, p. 1.

TypE.-Univ. of California Museum of Paleontology; a complete
shell.

TypE LoCALITY AND HeRIZON.-Barstow syncline, Mojave Desert, San
Bernardino County, California, U.S.A.; Barstovian faunal age, Late Mio-
cene.

GEOLOGIC RANGE.-Late Miocene.
GEOGRAPHIC RANGE.-Southeastern California, U.S.A.
REMARKS.-Close to G. brattstromi ( as G. depressus) and G. pansus

according to Brattstrom ( 1961).

Gopherus tneglectus C Brattstrom)

Gopherus neglectus Brattstrom 1961, p. 544, Bgs. 1-5.

TypE.-California Institute of Technology; an almost entire shell.
TypE LoCALAY AND H0RIZON.-Key Quarry, Ventura County, Cali-

fornia, U.S.A.; Upper Sespe Formation, Whitneyan faunal age, Late
Oligocene.

GEOLOGIC RANGE.-Late Oligocene.
GEOGRAPHIC RANGE.-California, U.S.A.
REMARKS.-Close to G. praeextans and G. laticuneus according to



186 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

Brattstrom ( 1961). G. neglectus and G. praeextans may be synonyms of
G. laticuneus.

Gopherus tpansus ( Hay)
Testudo pansa Hay 1908, p. 420, pl, 71, fgs. 1-2, text figs, 550-1,
Copherus pansa Williams 195Oa, p. 30.

TypE.-American Museum of Natural History; a complete shell and
pelvis.

TYPE LOCALITY AND HORIZON.-Near the state lines of Colorado and
Nebraska, north of Sterling, Weld County, Colorado, U.S.A.; Pawnee
Creek Formation, Barstovian faunal age, Late Miocene.

GEOLOGIC RANGE.-Late Miocene.

GEOGRAPHIC RANGE.-Southwestern Nebraska and northeastern Col-
orado, U.S.A.

REMARKS.-Close of G. brattstromi ( as depressus) and G. mohauetus,
according to Brattstrom ( 1961). The ending of the species name has
been changed to correspond with the masculine gender of Gopherus.

Gopherus tpertenuis ( Cope)
Testudo pertenuis Cope 1892b, p. 226.

TYPE.-Univ. of Texas; only a few fragments remain of the original,
almost complete type shell ( Wilson 1950).

TypE LoCALITY AND HORIZON.-Near Mt. Blanco, Crosby County,
Texas, U.S.A.; Early Blancan faunal age, Late Pliocene.

GEOLOGIC RANGE.-Late Pliocene.
GEOGRAPHIC RANGE.-Northwestern Texas, U.S.A.
REMARKS.-Probably very close to G. canyonensis and G. laticauda-

tus.

Gopherus polimhemus C Daudin)
Testudo polyphaemus Bartram 1791, p. 18 (nomen nudum).
Testudo polvphemus Daudin 1802.
Testudo depressa Guerin-Meneville 1829, p. 5, pl. 1, fig. 1.
Testudo gopher Gray 1844, p. 4.
Xerobates carolinus Agassiz 1857, p. 447.
Gopherus potyphemus Stejneger 1893, p. 161.
Gopherus carolinus Shaler 1888, p. 37.
Gopherus praecedens Hay 1916a, p. 55 ( Late Pleistocene, Vero, Florida).
Gopherus polt/phemus polvphemus Mertens and Wermuth 1955, p. 371.

TypE.-None designated.
TypE LOCALITY.-Savannah, Georgia, U.S.A. (by Schmidt 1953).
GEOLOGIC RANGE.-Late Pleistocene to Recent of Florida ( Hay 1930,

Holman  1958, 1959) and South Carolina. -



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 187

RECENT GEOGRAPHIC RANGE.-Southeastern United States ( Georgia,
Florida, and South Carolina west to the Mississippi River along the Gulf
Coast).

REMARKS.-Closely related to G. #avomarginatus C Auffenberg
1966b, in press). G. praecedens was originally believed to originate
from Early Pleistocene beds ( Hay 1916a), but these have since been
shown to be Late Pleist6cene and Recent in age ( Weigel 1962).

Gopherus tpraeextans ( Lambe)
Testudo praeextans Lambe 1913, p. 61.
Copherus praeextans Williams 195Oa, p. 29.

TypE.-Geological Survey ( Canada ); a shell.
TypE LOCALITY AND HORIZON.-Sage Creek, Niobrara County, Wy-

oming, U.S.A.; Orellan faunal age, Lower Brule age, Middle Oligocene.
GEOLOGIC RANGE.-Middle Oligocene.
GEOGRAPHIC RANGE.-Southeastern Wyoming, U.S.A.
REMARKS.-Probably only a chronologic race of G. laticuneus. Gil-

more ( 1946) described the osteology of an almost entire skeleton.

Gopherus f undabunus ( Loornis)
Testudo undabuna Loomis 1909, p . 25 .
Gopherus undabuna Williams 1950a, p. 30.
Geochelone undabuna Auffenberg 1964a, p. 6.

TYPE.-Amherst College; shell.
TYPE LOCALITY AND HORIZON.-Muddy Creek, Laramie County,

Wyoming, U.S.A.; Upper Harrison beds, Arikareean faunal age, Early
Miocene.

GEOLOGIC RANGE.-Early Miocene.
GEOGRAPHIC RANGE.-Eastern Wyoming, U.S.A.
REMARKS.-The ending of the species name has been changed to the

masculine gender of Gopherus. My earlier assignment to Geochelone is
here believed incorrect.

Genus Homopus Dumeril and Bibron
Lorica Walbaum 1782, p. 71.
Testudo Thunberg 1787 , p . 181 ( part ).
Chersine Merrem 1820, p. 30 ( part).
Homopus Dumeril and Bibron 1835, p. 145.
Chersobius Fitzinger 1835, p. 108 ( Testudo signata Schoepff).
Horopus Haycroft 1891, p. 341 ( typographical error).
Pseudomopus Hewitt 1931, p. 496 ( Testudo signata Schoepff).

GENOTYPE.-Homopus areolatus ( Thunberg).
DEFINITION.-Small Old World tortoises, with triturating surfaces of



188 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

maxilla and premaxilla without ridges; maxillary not entering roof of
palate; prootic ·narrowly exposed dorsally; quadrate enclosing stapes;
centrum of third cervical biconvex, no submarginal scutes; gulars divided;
gular region only slightly thickened.

GEOLOGIC RANGE.-No fossils known.
GEOGRAPHIC RANGE.-South Africa ( possibly into Southwest Africa).
REMARKS.-Contains four extant species: H. areolatus, H. boulengeri,

H. femoralis, and H. signatus. The most recent monographic treatment
is Loveridge and Williams ( 1957).

REFERENCES.-Skeleton: Boulenger 1889, 1890a, Auffenberg 1966b,
Gray 1873c, Williams 195Ob.

Genus Kinixys Bell
Kinixys Bell 1827, p. 398.
Cinothorax Fitzinger 1835, pp. 108, 111, 121.
Cir:irys Fitzinger 1843, p. 29.
Kinothorax Gray 1873a, p. 16.
Testudo Shaw 1802, p. 59 (part).
Homopus Lataste 1886, p. 286 ( part).
Malacochersus Loveridge 1942, p. 248 ( in error).

GENOTYPE.-Kinixys erosa ( Schweigger).
DEFINmoN.-No ridges on triturating surfaces of maxillary and pre-

maxillary; prootic well exposed dorsally; quadrate enclosing stapes;
third cervical biconvex; carapace of adult hinged; submarginal scutes
present; gular region greatly thickened and projected; gulars divided.

GEOLOGIC RANGE.-?Oligocene of France to Recent ( see Remarks).
GEOGRAPHIC RANGE.-Africa from 17° N., S. to Bechuanaland and

Natal, South Africa. May be introduced in Malagasy Republic.
REMARKS.-Three extant species are recognized: K. belliana, K.

erosa, K. homeana. Cinixys couzieri, Oligocene of France, is here ten-
tatively placed in Ptychogaster on the basis of basic shell morphology.

REFERENCES.-Skeleton: Gray 1873b, Boulenger 1889, Siebenrock
1910, Ruckes 1937, Williams 195Ob, Kilias 1957, Villiers 1958, Auffenberg
1966b.

Genus Malachochersus Lindholm
Cinicys Tornier 1896, p. 2 (part) [sicl (not of Gray).
Cinixys Procter 1922, p. 515.
Malacochersus Lindholm 1929, p. 285.
Testudo Siebenrock 1903a, p. 185.

GENOTYPE.-Testudo tornieri Siebenrock ( = Malacochersus torhieri
[Siebenrock]).

DEFINmON.-An African genus with a persistently fenestrated, very



1974 AUFFENBERG: - FOSSIL TORTOISE CHECKLIST 189

flat shell; a single suprapygal; gular region not thickened or projected;
gulars divided; triturating surface of maxilla strongly rigid; no median
premaxillary ridge; prootic well exposed dorsally; quadrate enclosing
stapes; submarginal scutes absent.

GEOLOGIC RANGE.-No fossils known.
GEOGRAPHIC RANGE.-Kenya, south to Tanganyika, Africa.
REMARKS.-The remark by Szalai ( 1934) that this genus possesses

teeth, and that it is thus similar to Stegochelys is incorrect. There is
only one extant species, M. tornieri.

REFERENCES.-Skeleton: Proctor 1922, Wettstein-Westersheim 1924,
Siebenrock 1904, Williams 195Ob, Richmond 1964.

Genus Psammobates Fitzinger

Testudo Linnaeus 1758, p. 199 ( part).
Chersine Merrem 1820, p. 32 ( part).
Psammobates Fitzinger 1835, pp. 108, 113, 122.
Psammobates Fitzinger 1843, p. 29 ( part).
Emys Gray 1844, p. 28 (part).
Clemmys Strauch 1862, p. 32 (part)
Peltastes Gray 1869, p. 173 (part)
Homopus Boettger, 1893, p. 8 ( part).
Chersinella Hewitt 1933, p. 259 (not of Gray).

GENOTYPE.-Testudo geometricus Linnaeus ( = Psammobates geo-
metricus [Linnaeus] ).

DEFINn'ION.z-Small South African tortoises with triturating surfaces
of maxilla not ridged; prootic narrowly exposed dorsally; quadrate en-
closing stapes; submarginal scutes absent; gulars divided; gular region
not greatly thickened or projected; anterior neurals hexagonal.

GEOLOGIC RANGE.-No fossils known.
~EOGRAPHIC RANGE.-Southern Africa.
REMARKs.-Of all the genera of tortoises, living and extinct, this one

has suffered the greatest vacillation in terminology. The most recent
species account is in Loveridge and Williams ( 1957). Three extant spe-
cies are recognized: P. geometricus, P. oculifer, P. tentorius.

REFERENCES.-Skeleton: Siebenrock 1897, Power 1932, Williams
195Ob, Auffenberg 1966b.

Genus Pyxis Bell
Pyxis Bell 1827, p. 395.
?Testudo Boulenger 1889, p . 45.
Bellemys Williams 195Ob, p. 553 ( in error).

GENOTYPE.-PYXis arachnoides Bell.
DEFINITION.-A monospeciEc genus of small tortoise from the Mala-



190 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

gasy Republic, characterized by a hinge at the anterior lobe of the
plastron.

GEOLOGIC RANGE.-No fossils known.
GEOGRAPHIC RANGE.-Malagasy Republic.

REMARKS.-Relationships not clear. Only one species recognized,
P. arachnoides.

REFERENCES.-Skeleton: Gray 18730, Williams 195Ob.

Genus iStylemys Leidy

Stylemys Leidy 195la, p. 173 ( not of Maack 1869).
Emys Leidy 1851a, p. 173.
Testudo Leidy 185lb, p. 173.
Geochelone Auffenberg 1963, p. 87 ( part).
Copherus Brattstrom 1961, p. 547 ( part).

GENOTYPE.-Stylemys nebrascensis Leidy.
DEFINITION.-An extinct Holarctic genus of tortoises characterized

by a premaxillary ridge; a symphyseal dentary groove; pleurals only
slightly alternately narrower and wider laterally; moderately long un-
specialized tail with some vertebrae lacking interpostzygopophyseal
notches; posterior epiplastral excavation shallower or absent; no hinge
on plastron or carapace; pleural bones not, or only slightly differen-
tiated; free proximal portion of ribs long; a single supracaudal scute;
nuchal scute longer than wide. The most recent description is that by
Auffenberg ( 1964d). Glaessner ( 1933) discussed the general problem
of "stylemiform" tortoises.

GEOLOGIC RANGE.-Eocene to Miocene of North America, Miocene
of Europe and Oligocene of Turkestan, U.S.S.R.

GEOGRAPHIC RANGE.-United States ( Oregon, California, Nebraska,
Colorado, South Dakota, Utah, Texas), France, and Turkestan.

REMARKS.-Shells described as Stylemys bottii from the Miocene of
France ( Stefano 1902a) and Stylemys karakolensis from the Oligocene of
Turkestan ( Riabinin 1927) may belong to this genus and are provision-
ally placed here.

Stylemys amphithorax ( Cope)

Testudo amphithorax Cope 1873a, p. 6.
St!/lemys amphithorax Auffenberg 1962d, p. 9.
Geochelone amphithorax Auffenberg 1963, p. 81 ( in error).

TypE.-American Museum of Natural History; cotypes, various parts
of three shells.

TypE LoCALITY AND HORIZON.-Head of Horsetail Creek, Weld
County, Colorado, U.S.A.; Horsetail Creek Member, White River Forma-
tion, Chadronian faunal age, Early Oligocene.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 191

GEOLOGIC RANGE.-Early Oligocene.
~EOGRAPHIC RANGE.-Northeastern Colorado and ( ?) South Dakota,

U.S.A.
REMARKS.-For description of skeleton see Auffenberg (1962d).

?Stylemys tbotti Stefano
?Stylemys bottii Stefano 1902a, p. 72.
St,lemys botti Williams 195Oa, p. 23.

TypE.-Presumably in the Museum of the School of Mines in Paris,
but not located on inquiry; most of carapace and plastron.

TYPE-LOCALITY AND HORIZON.-France; Miocene.
GEOLOGIC RANGE.-Miocene.
GEOGRAPHIC RANGE.-France.
REMARKS.-Generic assignment uncertain.

?St!/temys fcalaverensis Sinclair
?St!/lemys  calaverends Sinclair 1903, p. 243.

TypE.-Univ. of California Museum of Paleontology; a partial shell.
TypE LoCALITY AND HORIZON.-2 mi. S. of Vallecito, Calaveras Coun-

ty, California, U.S.A.; auriferous gravels, Miocene.
GEOLOGIC RANGE.-Miocene.
GEOGRAPHIC RANGE.-Northern California, U.S.A.
REMARKs.-Generic assignment not entirely clear because of frag-

mentary nature of the specimen.

Stylemys tcapax Hay
Stylemgs nebrascensis Cope 1885, p. 770 ("No. 2") (part)
St!/tem!/s cat,ax Hay 1908, p. 392, figs. 498-499.

TypE.-American Museum of Natural History; a complete shell.
TYPE LOCALITY AND HORIZON.-Near junction of North and South

Forks of John Day River, Grant County, Oregon, U.S.A.; Middle John
Day series, Arikareean faunal age, Early Miocene.

GEOLOGIC RANGE.-Early Miocene.
GEOGRAPHIC RANGE.-Central Oregon, U.S.A.

Stylemys t conspecta Hay
Stylemys nebrascensis Cope 1885, p. 769 (part).
Stylemys conspecta Hay 1908, p. 393.

TypE.-American Museum of Natural History; a shell.
TypE LoCALITY AND HORIZON.-Near junction of North and South



192 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

Forks of John Day River, Grant County, Oregon, U.S.A.; Middle John
Day series, Arikareean faunal age, Early Miocene.

GEOLOGIC RANGE.-Early Miocene.
GEOGRAPHIC RANGE.-Central Oregon, U.S.A.

Stylemys tkarakolensis Riabinin
St!/temus karakotensis Riabinin 1927, p. 193.
Stylemys" karakolensis Glaessner 1933, p. 360 [sic].

TypE.-Geological Institute, Academy of Sciences ( Leningrad,
U.S.S.R. ); a complete shell.

TYPE LOCALITY AND HORIZON.-Karakul, E. of Lake Issukkul, Sihk-
iang Province, Turkestan, U.S.S.R.; Oligocene ( or Early Miocene).

GEOLOGIC RANGE.-Oligocene (?).
GEOGRAPHIC RANGE.-Turkestan.

REMARKS.-Thought to be close to S. conspecta by Riabinin ( 1927).
Perhaps part of Geochelone, subgenus Indotestudo.

?Stylemys f ligonia ( Cope)
Testudo ligonius Cope 1873a, p. 6.
Testudo ligonia Cope 1873b, p. 19.
?Stylemys ligonia Auffenberg 1962d, p. 8 [sic].
Geochetone ligonia Auffenberg 1963, p. 92 ( in error).

TypE.-American Museum of Natural History; part of nuchal bone,
some peripherals and parts of the plastron.

TypE LOCALrrY AND H0RIZON.-Head of Horsetail Creek, Weld
County, Colorado, U.S.A.; Horsetail Creek Member, White River Forma-
tion, Chadronian faunal age, Early Oligocene.

GEOLOGIC RANGE.-Oligocene.
GEOGRAPHIC RANGE.-Northeastern Colorado, U.S.A.
REMARKS.-Probably conspecific with S. amphithorax ( see Auffen-

berg 1962d).

St!,lemys tnebrascensis Leidy

Emys hemispherica Leidy 185la, p. 173.
Emys hemispherica Leidy 185la, p. 173.
Testudo tata Leidy 185la, p. 173.
Emys oweni Leidy 185la, p. 327.
Emus culbertionii Leidy 1852a, p. 34.
Testudo hemispherica Leidy 1852a, p. 59.
Testudo nebrascensis Leidy 1852a, p. 59.
Testudo ott)eni Leidy 1852a, p. 59.
Testudo culbertsonii Leidy 1852a, p. 59.
Emys nebrascends Leidy 1852b, p. 65.
Emus lam Leidy 1852b, p. 65.
Stylemus culbertsonii Cope 1870, 8 124.



1974 AUFFENBERC: FOSSIL TORTOISE CHECKLIST 193

TYPE.-U. S. National Museum; most of the carapace and plastron.
TypE LoCALITY AND HORIZON.-South Dakota, U.S.A.; Brule series,

Orellan member, White River Formation, Orellan faunal age, Middle
Oligocene.

GEOLOGIC RANGE.-Middle Oligocene.
GEOGRAPHIC RANGE.-Nebraska, Colorado, Wyoming, North Dakota,

and South Dakota, U.S.A.
REFERENCES.-Skeleton. Case 1925, 1936, Auffenberg 1961, 1964d,

1966b, Hay 1907.

Stylemys tuintensis (Gilmore)
Testudo uintensis Gilmore 1915, p. 150.
?St!/temys uintensis Auffenberg 19624 p. 10.

TYPE.-Carnegie Museum; a complete shell.
TypE LoCALITY AND HomzoN.-South of Kennedy's Hole, 100 rods

west Dragon-Vernal Road, Uinta Basin, Uinta County, Utah, U.S.A.;
Horizon B or C., Uinta Formation, Uintan faunal age, Late Eocene.

GEOLOGIC RANGE.-Late Eocene.
GEOGRAPHIC RANGE.-Eastern Utah, U.S.A.
REMARKS.-A primitive member of the genus, little differentiated

from Geochelone.

Genus Testudo Linnaeus
Testudo Linnaeus 1758, p. 197.
Chersine Merrem 1820, p. 43 ( Type: T. graeca).
Emys Meyer 1834, p. 121 (part)
Chersus Brandt 1852, p. 331 ( part).
Homopus Blyth 1854, p. 642 ( part) ( Type: H. burnedi).
Peltastes Gray 1869, p. 171 (Type: T. graeca L.).
Chersinetta Gray 187Ob, p. 725 ( Type: T. gmeca L.).
Testudinelta Gray 187Ob, p. 658 ( Type: T. homieldii Gray ).
Peltonia Gray 1872a, p. 4 ( Type: T. graeca L.).
Medaestia Wussow 1916, p. 170 ( Type: T. graeca L. ).
St!/lemvs Auffenberg 1964d, p. 121 ( part).
Agrionemys Khozatsky and Mlynarski 1966, p. 123 ( part) ( Type: T. horsjie~di Gray ).
Agrione Khozatsky and Mlynarski 1966, p. 123 ( typographical error).
Protestudo Chkikvadze 1970c, p. 7 ( part) ( Type: T. bessarabica).

Gif:NOTYPE.-Testudo graeca Linnaeus.
DEFINITION.-A Eurasian and North African genus of tortoises char-

acterized by the presence of a nuchal scale; suprapygal single or divided
and, if divided, a larger anterior one embracing a smaller posterior scale;
posterior lobe of plastron more or less movable in one or both sexes.
Three subgenera are recognized here: Pseudotestudo, Agrionemys, and
Testudo.



194 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

GEOLOGIC RANGE.-Miocene to Recent.
GEOGRAPHIC RANGE.-Recent forms are distributed in Southern

Europe, western and south central Asia, and northern Africa. Fossils
are found over much of the central and southern part of the Palearctic.

REMARKS.-Protestudo of Chkikvadze ( 1970c) is here considered a
synonym of Testudo because the generic characters given are not suffi-
ciently distinct in view of those for the remaining testudinid taxa.

Subgenus Agrionemys Khozatsky and Mlynarski
Testudo Pallas 1811, p. 19 (part).
Chersus Brandt 1852, p. 331 ( part).
Homopus Blyth 1854, p. 642 ( Type: H. burnesii).
Testudinella Gray 187Ob, p. 658.
Agrionemys Khozatsky and Mlynarski 1966, p. 123 ( as genus).
Agrione Khozatsky and Mlynarski 1966, p. 123 ( typographical error).

TYPE SPECIES.-Testudo hors/ieldi Gray.
DEFINITIoN.-A subtropical Asian subdivision of the genus Testudo

characterized by moderately ridged maxillary; four toes on the anterior
limbs; humero-pectoral scute usually crossing posterior part of the ento-
plastron; and two suprapygal bones in which the upper embraces the
lower.

GEOLOGIC RANGE.-Recent only at this writing, but undoubtedly
with many Pliocene and Pleistocene members in Asia.

GEOGRAPHIC RANGE.-Turkestan to northwestern West Pakistan.
REMARKS.-Apparently a primitive subgroup of the genus Testudo.

It may eventually be found that many of the small globular Tertiary and
Pleistocene tortoises of Southern Russia and China belong to this sub-
genus.

Testudo ( Testudo) hors#eldi Gray

Testudo geometrica Pallas 1811, p. 19 ( part).
Testudo horsfieldii Gray 1844, p. 74 ( Afghanistan).
Chersus iberus Brandt 1852, p. 331 ( part).
Homopus bumesii Blyth 1854, p. 642 ( Afghanistan).
Testudinella hors~eldii Gray 187Ob, p. 658.
Homopus horsfieldi Syevertzov 1873, p. 71.
Testudo baluchiorum Annandale 1906, pp. 75, 205, pl. 2, fig. 1 ( Baluchistan).
Agrionemys horsfieldi Khozatsky and Mlynarski 1966, p. 123.

TypE.-British Museum ( Natural History) ; mounted specimen.
TypE LOCALITY.-Afghanistan.
GEOLOGIC RANGE.-NO fossils known, but included here because

some of Plio-Pleistocene fossils will certainly be referred here with fur-
ther study.

GEOGRAPHIC RANGE.-Eastern shore of the Caspian Sea, Turkestan



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 195

to Aral 'Skoye More' ( Aral Sea), Iran, Afghanistan, and northwestern
West Pakistan.

REMARKS.-The report of a Nepalese specimen ( Giinther 1864) and
verbal accounts of tortoises in the Dera-Dun area of India that seem to
represent this tortoise suggest hors#eldi will eventually be found to in-
habit much of the foothill area of the Himalaya Mountains. The recent
erection of Agrion€mys for reception of hon#eldi ( Khozatsky and Mly-
narski 1966) is not followed here because I consider the purported
generic characters within the generic spectrum of variability of Testudo.

REFERENCES.-Morphology. Skeleton: Auffenberg 1966b, Williams
1950b, Mlynarski 1966b, Khozatsky and Mlynarski 1966. Serology. Obst
and Ambrosius 1971.

Subgenus Pseudotestudo Loveridge and Williams

Testudo Lichtenstein 1823, p. 91 ( part).
?Chersus Fitzinger 1955, p. 252 ( part).
Peltastes Gray 187Ob, p. 657 ( part).
Pseudotestudo Loveridge and Williams 1957, p. 276 ( as subgenus).

TYPE SPECIES.-Testudo ( Pseudotestudo ) kleinmanni Lartet.
DEFINITION.-An African subdivision of the genus Testudo charac-

terized by usually having a single supracaudal; maxillary without ridges;
larger exposed anterior palatine foramina; quadrate not enclosing stapes;
plastron of all adults with clearly movable posterior lobe.

GEOLOGIC RANGE.-NO fossils known.
GEOGRAPHIC RANGE.-WeStern Cyrenaica, Libya east to northern

United Arab Republic and northern Sinai.
REMARKS.-Composed of only one species, T. kleinmanni, which is

close to the gra€ca group according to Loveridge and Williams ( 1957 ).
REFERENCES.-Skeleton: Siebenrock 1906, Williams 195Ob, Love-

ridge and Williams 1957.

Subgenus Testudo Linnaeus
Testudo Linnaeus 1758, p . 198.
Emys Meyer 1834, p. 121 (part).
Peltastes Gray 1870c, p. 11 ( part).
Chersinella Gray 1873c, p. 725, pl. 60, fg. 4 (part)
Stulem!/s Auffenberg 1964d, p. 121 ( part).

TYPE SPECms.-Testudo graeca Linnaeus ( = Testudo [Testudo]
graeca Linnaeus).

DEFINITION.-A Eutasian subdivision of the genus Testudo character-
ized by a weakly or moderately ridged maxillary; anterior palatine fora-
men, partially concealed in ventral view; quadrate enclosing stapes;



196 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

plastron of adults with more or less movable posterior lobe in one or
both sexes.

GEOLOGIC RANGE.-Miocene to Recent.
GEOGRAPHIC RANGE.-Northern Africa, southern Europe, and western

Asia.
REMARKS.-Contains three extant species, T. marginata, T. her-

manni, and T. graeca, in addition to the many fossil forms.
REFERENCES.-Morphology. General Anatomy: Thompson 1932;

Skeleton: Simonetta 1960.

Testudo talba ( Chkikvadze)
Protestudo alba Chkikvadze 1971, p. 245, Bg. 1.

TypE.-Georgian Academy of Science ( Tbilisi); right epiplastron.
TypE LOCALITY AND HORIZON.-Georgian S.S.R., eastern Kazakhstan,

Zaisan Valley; Upper Oligocene to Lower Miocene.
GEOLOGIC RANGE.-Upper Oligocene to Lower Miocene.
GEOGRAPHIC RANGE.-Georgian S,S.R.
REMARKS.-Originally placed in the genus Protestudo, but here con-

sidered a synonym of Testudo.

Testudo Crestudo ) tantiqua Bronn
Testudo antiqua Bronn 1831, p. 215.
Emys striata Meyer 1834, p. 121.
Testudo striata Giebel 1847, p. 58.
Emys mellingi Peters 1869, p. 122.
Testudo amiatae Pantanelli 1893, p. 129 [sic] ( Vindobonian Miocene, Italy).
Testudo noviciensis ( Nouel MS) Deperet 1895, p. 412 ( Burdagalian Miocene,

Austria).
Testudo amiatae Reinach 1900, p. 17 ( part).
Testudo promarginata Reinach 1900, p. 74 ( part).
?Testudo neouiciensis Koch 1904, p. 60 ( part) ( typographical error).
Testudo antiqua var. nouiciensis Glaessner 1933, p. 364.
Testudo antiqua nouiciensis Mlynarski 1955a, p. 173.
Testudo praeceps Haberlandt 1876, p. 243 ( Vindobonian Miocene, Austria).
Testudo kalksburgensis Toula 1896, p. 915 ( Vindobonian Miocene, Austria).
Testudo leithi Siebenrock 1915, p. 84 ( part).
Testudo antiqua var. praeceps Glaessner 1933, p. 362.
Testudo antiqua pmeceps Glaessner 1935, p. 125.
Testudo craverii Portis 1879, p. 121 [sic] ( Sarmatian Mi6cene, Italy).
Testudo craveri Pantanelli 1893, p . 135.
Testudo onward Teppner 1913, p. 384 ( typographical error).
Testudo claweri Riabinin 1915, p. 11 ( typographical error).
Testudo graweri Bataller 1926, p. 149 ( typographical error).
Testudo crawenii Riabinin 1926, p. 60 ( typographical error).
Testudo escheri Pictet and Humbert 1856, p. 17 [sic] ( Vindobonian Miocene,

Switzerland).
Testudo minuta Fraas 1870, p. 51 ( in error).



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 197

Testudo kalksbergensis var. steinheimensis Straesche 1931, p. 4 ( Vindobonian Mio-
cene, Germany).

Testudo graeca Koch 1902, p. 311 (part).
Testudo syrmiensis Koch 1904, p. 61 ( Pontian Pliocene, Hungary).
Testudo canetotiana Lartet 1851, p. 38 ( Vindobonian Miocene, France).
Testudo cannetotiana Szalai 1935, p. 380 ( typographical error).
Stul€mys canetotiana Auffenberg 1964d, p. 121.
Testudo csakcarensis Szalai 1934, p. 119 ( Sarmatien Miocene, France).
Testudo csakuariensis Mlynarski 1966a ( typographical error).

TypE.-Fursten von Furstenberg? collection; a shell.
TYPE LOCALITY AND HoRIZON.-Hohenh6ven, Engen in Hegau, Ger-

many; Late · Miocene.
GEOLOGIC RANGE.-Early Miocene to Early Pliocene.
GEOGRAPHIC RANGE.-Early Miocene of Austria. Late Miocene of

France, Germany, Austria, Italy, Greece, and Switzerland. Early Plio-
cene of Poland ( Mlynarski 1962) and Hungary ( Mlynarski 1955a).

REMARKS.-Good descriptions are found in Campana ( 1917a) of
specimens referred to T. amiatae from Tuscany, Italy, and in Meyer
( 1867) of the type.

Not all workers will agree with my disposition of most species placed
in the synonomy of T. antiqua. The principal philosophy behind this
action is outlined in a series of often confusing statements expressing
the close relationship and possible conspecificity of these forms in one
combination or another. A comparison of conclusions by Roger 1902,
Stefano 1902b, Siebenrock 1915, Glaessner 1933, 1935,· Szalai 1934, 1935,
Thenius 1951, and Mlynarski 1955a, 1956, 1962, regarding the relation-
ships of these forms ( often named on the most meager material) should
prove the practicality of the present action.

Siebenrock ( 1915) and Mlynarski ( 1955a) consider Testudo prae-
ceps Haberlandt to be quite distinct from antique and the latter states
it may be close to Stylemys. Thenius ( 1952) considers the "stylemys"
characters of praeceps as an individual variation of antiqua, with which
I concur.

Testudo ( Testudo) tbosporica Riabinin
Testudo bosporica Riabinin 1945, p. 127.

TypE.-Paleontological Museum ( Leningrad ); a poor shell.
TYPE LOCALITY AND HORIZON.-Mt. Opouk, near Kertch, Crimea,

U.S.S.R.; Sarmatian faunal age, Late Miocene.
GEOLOGIC RANGE.-Late Miocene.
GEOGRAPHIC R~NGE.-Crimea.
REMARKS.-Close to T. hermanni or T. graeca according to Riabinin

( 1945); in the antiqua-graeca phyletic line according to Khozatsky
(1947).



198 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

Testudo (Testudo) fcatalaunica Bataller
Testudo catalaunica Bataller 1926, p. 149.

TYPE.-Seminario Councilar de Barcelona ( Spain ); a shell.
TYPE LOCALITY AND HORIZON.-St. Quirce, near Bafcelona, Barcelona

Province, Catalonia region, Spain; Vindobonian Miocene.
GEOLOGIC RANGE.-Middle Miocene.
GEOGRAPHIC RANGE.-Northeastern Spain.
REMARKS.-Relationship close to graeca group and probably a syn-

onym of T. antiqua. T. catalaunica var. irregularis Bergounioux ( 1958)
is intended as a niorphotype, not a subspecies, and therefore has no
validity as a trinomial. It is from the same locality as the type.

Testudo ( Testudo) f cllonica Bergounioux

Testudo celonica Bergounioux 1958, p. 171; fgs. 13, 14; pls. 30,31.

TypE.-Seminario Councilar de Barcelona ( Spain ); a partial shell.
TYPE LOCALITY AND HORIZON.-San Celoni, near Barcelona, Cata-

lonia region, Spain; Vellesian Miocene.
GEOLOGIC IUNCE.-Late Miocene.
GEOGRAPHIC RANGE.-Northeastern Spain.
REMARKS.-Probably a synonym of T. antiqua.

Testudo ( Testudo) f cernovi Khozatsky

Testudo cemoui Khozatsky 1948, p. 92.

TypE.-Zoology Institute, Academy of Sciences ( Ukranian S.S.R.);
anterior parts of carapace and plastron.

TypE LoCALITY AND HORIZON.-Village Gol'ma, Baltsk region,
Ukrainian S.S.R.; Kucurgan deposits, Late Pliocene.

GEOLOGIC RANGE.-Pliocene.
GEOGRAPHIC RANGE.-Ukraine.

REMARKS.-CloSe to the antiqua-graeca line, according to Mlynarski
( 1955a), and particularly to T. graeca and T. hermanni C Mlynarski
1968).

Testudo (Testudo) tcorrogi Bergounioux

Testudo corrot/i Bergounioux 1933a, p. 508.

TypE.-Geologic Laboratory, Faculty of Sciences ( Marseille,
France ); a carapace.

TYPE LOCALITY AND HORIZON.-Lutetian Basin of Palette, Aix-en-

Provence, Bouches-du-Rhone Dept., France; Lutetian faunal age, Middle
Eocene.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 199

GEOLOGIC RANGE.-Middle Eocene.
GEOGRAPHIC R~NGE.-France.
REMARKS.-Related to the graeca group according to Bergounioux

( 1933a) and near base of the genus Testudo. Discussed further by Ber-
gounioux ( 1935).

Testudo tdarewskii ( Chkikvadze)
Protestudo darewskii Chkikvadze 1971, p. 46, fig. 2.

TypE.-Georgian Academy of Science ( Tbilisi); right epiplastron.
TypE LOCALITY AND HORIZON.-Georgian S.S.R., eastern Kazakhstan,

Zaisan Valley; Middle to Upper Miocene.
GEOLOGIC RANGE.-Middle to Upper Miocene.
GEOGRAPHIC RANGE.-Georgian S.S.R.
REMARKS.-Originally placed in the genus Protestudo, but is here

considered a synonym of Testudo. T. darewskii is probably close to
T. alba.

Testudo ( Testudo) tdenizoti Bergounioux
Testudo denizoti Bergounioux 1935, p. 87-, pl. 5, fg. 1.
Testudo denizotti Mlynarski 1955a, p. 49 ( typographical error in English version only)

TypE.-Paleontological Laboratory of Geology, Faculty of Sciences
( Marseille ); a carapace.

TYPE LOCALITY AND HORIZON.-Near Puy Laurens, Tarn Department,
France; Stampian faunal age, Middle Oligocene.

GEOLOGIC RANGE.-Middle Oligocene.
GEOGRAPHIC RANGE.-France.
REMARKS.-Close to T. catalaunica according to Bergounioux ( 1935).

In antiqua-graeca phyletic line according to Mlynarski ( 1955a), Ber-
gounioux ( 1935), and Bram ( 1951).

Testudo ( Testudo ) fdoduni Gray
Testudo doduni Gray 183lb, p. 47.

TypE.-Location unknown to author; fragments of shell.
TYPE LOCALITY AND HORIZON.-Castelnaudary, Aude Department,

France; Lutetian faunal age, Middle Eocene.
GEOLOGIC RANGE.-Eocene.
GEOGRAPHIC RANGE.-France.
REMARKS.-Probably indeterminate.

Testudo (Testudo) f globosa Portis
Testudo globosa Portis 1890, p. 3.



200 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

TypE.-Firenze Museum; a shell.
TypE LOCALITY AND HORIZON.-Valley of the Amo River, Tuscany,

Italy; Villafranchian faunal age, Early Pleistocene.
GEOLOGIC RANGE.-Early Pleistocene.
GEOGRAPHIC RANGE.-Northern Italy.
REMARKS.-Presumed to be close to the graeca-hermanni line by

Portis ( 1890), closest to T. hermanni (Mlynarski 1962 ). T. szalaii may
be a synonym of T. globosa. In antiqua-graeca phyletic line and perhaps
a synonym of T. antiqua ( Glaessner 1933).

Testudo ( Testudo) graeca Linnaeus

Testudo graeca Linnaeus 1758, p. 198.
Testudo pusilla Linnaeus 1766, p. 353 ( part).
Testudo terrestris Forskal 1775, p. 12 ( -Testudo [Testudol graeca terrastris Forskal,

Lebanon Mts., Israel, by Wermuth 1958) ( not Testudo terrestris of Fermin
1765).

Testudo ibera Pallas 1814, p. 18 ( -Testudo [Testudo] graeca ibm Pallas, Middle
Kura-Tales in Caucasus by Mertens 1946).

Testudo georgicana Guldenstedt 1814 (In Pallas) ( nomen nudum)
Testudo ecaudata Pallas 1827, p. 19 ( Tiflis, Iran at the Caspian Sea).
Testudo geometrica Hohenacker 1831, p. 364 ( Caucasus) ( part).
Testudo zolkafa Forskal 18312, p, 13, In Gray· ( Arabia) ( nomen nudum).
Testudo zohalfa Forskal 1835, p. 44, In Dumeril and Bibron ( Arabia) ( nomen

nudum).
Testudo mauritanica Dumeril and Bibron 1935, p. 44 ( part).
Testudo mauritonica Kercado 1835, p. 35 ( typographical error).
Testudo marginata Gervais 1836, 8 309 (not of Schoepff).
Testudo ibera Gervais 1836, p. 309 (not of Pallas).
Testudo whitei Bennett 1840, p. 361 ( In White).
Testudo graeca var. mauritanica Schlegel 1841, p. 106.
Testudo pusilla Strauch 1862, p. 67 ( Transcaucasia) ( part).
Pettastes marginatus var. whitel Gray 1870c, p. 11.
Chersinella graeca Gray 1873b, p. 725, pl. 60, fig. 4
Testudo zarudyni Nikolsky 1869, p, 369 ( = Testudo [Testudo] graeca zarudnt/i

Nikolsky, Birdschau Prov., Eastern Iran, by Mertens 1946).

TypE.-Location unknown to author.
TypE LoCALITY.-Santa Cruz, Oran, Algeria, Africa (by Mertens and

Muller 1928 ).
GEOLOGIC RANGE.-Pleistocene of Motocco ( Lecointre 1926), France

(Henri-Martin 1946) and Recent.
GEOGRAPHIC RANGE.-Northwestern Africa from Morocco. and Malta

to Cyrenaica region of Libya.
REMARKS.-~ee Mertens ( 1946) for monographic treatment, and

Obst and Ambrosius ( 1971) for a recent discussion of relationships to
European members of the genus.

REFERENCES.-Morphology. Tympanum: Filogamo 1849; Bone His-

tology: Amprino and Godina 1947; Skull: Gray 1873c, Siebenrock 1897,



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 201

Bellairs 1949, Staesche 1961; Myology: Favaro 1903, Hacher and Schu-
macher 1955, Stammer 1959; Hyobranchial Apparatus: Siebenrock 1898,
Bender 1914; Skeleton: Lydekker 1889a, Briihl 1896, Siebenrock 1910,
Nopsca 1926a, 1926b, Mertens 1936, Khozatsky 1941, Bannikov 1947,
Williams 195Ob, Loveridge and Williams 1957, Staesche 1961, Auffen-
berg 1966b; Serology: Obst and Ambrosius 1971.

Testudo ( Testudo) hermanni Gmelin
Testudo hermanni Gmelin 1788, p. 1041 ( unknown).
?Testudo ( Emys) tcanstadiensis Plieninger 1847, p. 208 ( Sinterkalk of Wurttemberg).
Testudo graeca bettai Lataste 1881, p. 396 ( unknown).
Testudo graeca Boulenger 1889, p . 177 ( part ).
Testudo graeca var. boettgeri Mojsisovics 1888, p. 242 ( not T. boettgeri Siebenrock)

( Orsova, Banat).
Testudo graeca var. hercegovinensis Werner 1899, p, 818 ( Trebinje, Hercegovina).
Testudo entiquest Parenzan 1932, p. 1160 ( Conca di Elbasson, Skumbi, Central

Albania).
Testudo hermanni hermanni Wermuth 1952, p. 161.

TypE.-None designated originally ( Wermuth 1952).
TypE LOCALITY AND HORIZON.-None designated.
GEOLOGIC RANGE.-Pleistocene of Italy ( Campana 1917b) and Wurt-

temberg ( Plieninger 1847) to Recent.
GEOGRAPHIC RANGE.-Southern Italy, Sicily; the Balkans south of the

Danube River ( except in Dobrua), to the Peloponnesus in Greece, north
of the Danube River only in southwestern Romania. Intergrades with
T. h. robertmertensi on Corsica, Sardinia, and in (?) northern Italy.

REMARKS.-In antiqua-graeca phyletic line according to Glaessner
(1933).

REFERENCES.-Morphology. Skeleton: Siebenrock 1910, Loveridge
and Williams 1957, Auffenberg 1966b; Shell: Staesche 1961; Serology:
Obst and Ambrosius 1971.

Testudo ( Testudo) thipparionum Wiman

Testudo hipparionum Wiman 1930, p. 41, pl. 6, fig. 2-2b.
Geochetone hipparionum Mlynarski 1968, p. 95.

TypE.-Institute of Vertebrate Paleontology and Paleonanthropology
( Peking ); a shell.

TYPE LOCALITY AND HORIZON.-Wuhsiang-Hsien, Shansi Province,
China; Pontian faunal age, Early Pliocene.

GEOLOGIC RANGE.-Early Pliocene.
GEOGRAPHIC RANGE.-Northern China.
REMARKS.-Represented in the Shansi Steppe fauna. Yeh ( 1963a)

described and figured a complete specimen referred to this species. Very



202 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

poorly defined by Wiman ( 1930) and undoubtedly conspecific with sev-
eral of his other species from the same horizon and area. I see no mor-
phological basis for including the species in Geochelone.

Testudo (Testudo) thonanensis Wiman
Testudo honanensis Wiman 1930, p, 43.
Testudo hannonensis Mlynarski 1955b, p. 170 ( typographical error).

TypE.-Institute of Vertebrate Paleontology and Paleoanthropology
( Peking ); a shell.

TypE LocALITY AND HORIZON.-Hsin-An-Hsien, Honan, China; Pon-
tian faunal age, Early Pliocene.

GEOLOGIC RANGE.-Early Pliocene and ?Miocene.
GEOGRAPHIC RANGE.-Honan, Shansi, and Kansu Provinces, China.
REMARKS.-In antiqua-graeca phyletic line according to Mlynarski

( 1955b). Undoubtedly conspecific with other described species from
the same horizon, and area.

Testudo ( Testudo) thypercostata Wiman

Testudo hypercostata Wiman 1930, p. 35, pl. 5, fig. 3-36.

TYPE.-Institute of Vertebrate Paleontology and Paleoanthropology
( Peking ); a shell.

TYPE LOCALITY AND HORIZON.-Ho Ch'u-Hsien, Shansi Province,
China; Pontian faunal age, Early Pliocene.

GEOLOGIC RANGE.-Early Pliocene.
GEOGRAPHIC RANGE.-Shansi, China.
REMARKS.-A very poorly defined species, probably an individual

variant and quite likely conspecific with T. shensiensis.

Testudo tilliberalis ( Chkikvadze)
Protestudo illiberatis Chkikvadze 1971, p. 246, fg. 3.

TypE.-Georgian Academy of Science ( Tbilisi); nearly complete
plastron imbedded in nodule.

TypE LoCALrrY AND HORIZON.-Georgian S.S.R., eastern Kazakhstan,
Zaisan Valley; Lower to Middle Pliocene.

GEOLOGIC RANGE.-Lower to Middle Pliocene.
GEOGRAPHIC RANGE.-Georgian S.S.R.
REMARKS.-Perhaps close to Testudo hors/ieldi.

Testudo ( Testudo) tkegenika Khozatsky

Testudo kegenika Khozatsky 1955, p. 51.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 203

TypE.-Geological Museum, Academy of Sciences ( U.S.S.R. ); a shell.
TypE LOCALITY AND HORIZON.-Upper Course Kegen River, Chula-

dir Ridge, Kazakhstan, S.S.R., U.S.S.R., Alura-Atiau Province, Kegen
Region, Tienshan "Neogene" ( = Upper Miocene?).

GEOLOGIC RANGE.-"Neogene" ( - Upper Miocene?).
GEOGRAPHIC R~NCE.-Kazakhstan, S.S.R.
REMARKS.-The age of this fossil tortoise is questionable ( Bazhanov

and Pigulevsky 1955). Mlynarski ( 1968) places this species in the genus
Geochelone without comment. Its characters are such that I prefer to
leave it in Testudo until proof of another assignment is forthcoming.

Testudo (Testudo) tkucurganica Khozatsky

Testudo kucurganica Khozatsky 1948, p. 94.

TYPE.-Zoology Institute, Academy of Sciences ( Ukrainian S.S.R.);
comprised of numerous fragments of shell from 40-50 specimens of dif-
ferent sizes.

TYPE LOCALrrY AND HORIZON.Novopetrovka, Alekseeva and other
localifies in the valley of the Kucurgan River, District Odessa, Ukrainian
S.S.R.; Kucurgan deposits, Late Pliocene.

GEOLOGIC RANGE.-Pliocene.
GEOGRAPHIC RANGE.-Ukraine.

REMARKS.-Close to the antiqua-graeca group according to Mlynar-
ski ( 1955a). Close to T. graeca and T. hermanni according to Khozatsky
(1948).

Testudo ( Testudo) f lamanoni Cuvier

?Testudo sp. Lamanon 1780, p. 868.
Testudo lamanonii Cuvier 1836, p. 486.
Testudo lamanoni Gervais 1859, p. 438.
Testudo lamanonis Bergounioux 1933a, p. 515 ( typographical error).

TypE.-Museum of Natural History ( Paris); an internal mold of a
carapace.

TypE LOCALITY AND HORIZON.-Near Aix-en-Provence, Bouches du
Rhone Department, Provence region, France; Oligocene.

GEOLOGIC RANGE.-Oligocene.
GEOGRAPHIC RANGE.-Provence, France.
REMARKS.-In antiqua-graeca phyletic line according to Mlynarski

( 1956). According to Bergounioux ( 1932) the specimen of this species
described by Gervais ( 1859) belongs to Clemmys uidali.

Testudo ( Testudo) flunellensis Almera and Bofill

Testudo luneHensis Almera and Bofill 1903, p. 106 [sic].
Testudo lunellensis Deperet 1906, p. 12.



204 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

TypE.-Museo Municipales Barcelona; a shell.
TypE LoCALITY AND HoRIzoN.-Cavern in Park Guell, Barcelona,

Spain; Early Pleistocene.
GEOLOGIC RANGE.-Pleistocene.
GEOGRAPHIC RANGE.-Near Barcelona, Spain.
REMARKS.-In the antiqua-graeca phyletic line according to Ber-

gounioux ( 1935). T. lunellensis var. iberica Bergounioux ( 1958) is in-
tended as the designation of a morphotype, not a subspecies, and there-
fore has no validity as a trinomial.

Testudo ( Testudo) macarovicii Mlynarski
Testudo pfaegraeca ibera Marcarovici and Vancea 1960, p. 377 ( nomen illicit.).
Testudo macarovicii Mlynarski 1969b, p. 152 ( new name).

TypE.-Designated by Mlynarski ( 1969b) as the many isolated frag-
ments of the carapace and plastron in the Geological Institute, A. I. Cuza
University ( Iasi, Romania).

TYPE Locurry AND HoRIZON.-Malusteni, Moldavia, Romania; As-
tian faunal age, Late Pliocene?

GEOLOGIC RANGE.-Late Pliocene?
GEOGRAPHIC RANGE.-Besides the type locality, known from Mana-

stirea, Minzatesti, Plesea, Beresti, Pruth, Roscani, and a number of other
localities, all in Moldavia, Romania.

REMARKS.-As Mlynarski ( 1969b) states this species is extremely
close to T. graeca ibera. It is indeed a pity that he formally renamed
Macarovici and Vancea's Testudo praegraeca ibera, when the illegality
of their name (Art. 52 and 57, Int. Code Zool. Nomen.) would have
afforded a convenient method to place it where it probably belongs-a
synonym of T. graeca.

Testudo ( Testudo ) marginata Schoepff
Testudo tabulkita var. campanutata Walbaum 1782, p. 124 ( nomen illegitum).
Testudo marginata Schoepff 1792, p. 58, pls. 11-12, fig. 1 ( unknown).
Testudo grafa Hermann 1804, p. 219 ( Greece, by Mertens and Wermuth, 1955).
Chersine marginata Merrem 1820, p. 31.
Chersine marginatus Wagler 1833, p. 138.
Testudo campanulata Strauch 1862, p. 65 ( Greece).
Pettastes marginatus Gray 1869, p. 173.
Testudo nemoratis Schreiber 1875, p. 557 ( Greece).

TypE.-Location unknown to author.
TYPE LOCALrry-Unknown.
GEOLOGIC RANGE.-No fossils known but included here because of

my conviction that further study of European fossils of Testudo will in·
dicate its presence.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 205

GEOLOGIC RANGE.-Southern Greece, north to Olymp, the island of
Skiros, southern Albania, and (?) Sardinia.

REMARKS.-Considered very close to T. graeca by Siebenrock ( 1910)
and placed in antiqua-graeca phyletic line by Glaessner ( 1933). Serologi-
cal analysis ( Obst and Ambrosius 1971 ) suggests a somewhat more dis-
tant relationship to graeca and hermanni.

REFERENCES.-Morphology. Skeleton: Siebenrock 1910, Williams
195Ob, Auffenberg 1966b; Hyobranchial Apparatus: Siebenrock 1898;
Skull: Siebenrock 1897; Serology: Obst and Ambrosius 1971.

Testudo ( Testudo) tmarmorum Gaudry
Testudo ma,momm Gaudry 1862, p. 502.
Testudo ( Chersis) marmorum Hoernes 1892, p. 245.
Testudo marmorem Nafiz and Malik 1933, p. 120 (typographical error).

TYPE.-Paris Museum; almost complete shell.
TypE LOCALITY AND HORIZON.-Pikermi, Greece; Pontian faunal age,

Early Pliocene.
GEOLOGIC RANGE.-Early Pliocene.
GEOGRAPHIC RANGE.-Greece and adjacent Turkey.
REMARKS.-Posterior part of plastron movable. Best description in

Gaudry ( 1862-7). Nuchal scute very narrow. Perhaps in antiqua-graeca
line, though Mlynarski ( 1955a) and Glaessner ( 1933) think not.

Testudo ( Testudo) toriens Portis

Testudo oriens Portis 1890, p. 9.

TypE.-Location unknown to author; a shell.
TypE LOCALITY AND HORIZON.-Valley Of the Arno River, Tuscany,

Italy; Villafranchian faunal age, Early Pleistocene.
GEOLOGIC RANGE.-Early Pleistocene.
GEOGRAPHIC RANGE.-Italy,

Testudo ( Testudo) tpusilla Bergounioux
Testudo pusilla Bergounioux 1936a, p. 21 ( not T. pusilla Linn.)

TypE.-Location unknown to author; partial shell.
TYPE LOCALrry AND HoRIZON.-Near Marseilles, Bouches du Rhone

Department, France; Stampian faunal age, Middle Oligocene.
GEOLOGIC RANGE.-Middle Oligocene.
GEOGRAPHIC RANGE.-France.
REMARKS.-Close to Geochelone nana according to Bergounioux

( 1936a), but retained in Testudo here on basis of available material.



206 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

Testudo ( Testudo) troguesi Bergounioux

Testudo rougesi Bergounioux 1936b, p. 58.

TYPE.-Geological Laboratory ( Clermont-Feuant, France); a shell.
TypE LOCALITY AND HoRIzoN.-Bard near Brioude, Haute Loire De-

partment, France; Oligocene.
GEOLOGIC RANGE.-Oligocene.
GEOGRAPHIC RANGE.-France.
REMARKS.-C1OSe to Testudo antiqua according to Bergounioux

(1936b).

Testudo ( Testudo) fsemenensis Bergounioux

Testudo semenensis Bergounioux 1955, pp. 145-152.

TypE.-Location unknown to author; a carapace.
TYPE LoCALITY AND HORIZON.-Djeber Semene, 110 km SE of Tunis,

Tunisia; Pontian faunal age, Early Pliocene.
GEOLOGIC RANGE.-Early Pliocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-Probably in antiqua„graeca line.

Testudo ( Testudo) fseminota Portis

Testudo seminota Portis 1890, p. 10, pl. 2, fig. 13,

TypE.-Location unknown to author; a shell.
TypE LOCALITY AND HORIZON,-Valley Of the Arno River, Tuscany,

Italy; Villafranchian faunal age, Early Pleistocene.
GEOLOGIC RANGE.-Early Pleistocene.
GEOGRAPHIC RANGE.-Italy.
REMARKS.-ClOSe tO hermanni-graeca.

Testudo ( Testudo ) iserresi Pictet

Testudo serresii Pictet 1845, p. 20.
Testudo serresi Bergounioux 1938a, p. 279.

TypE.-Location unknown to author; a shell.
TypE LoCALrrY AND HoRIZON.-Montpellier, Herault Department,

France; Astian faunal age, Late Pliocene.
GEOLOGIC RANGE.-Late Pliocene.
GEOGRAPHIC RANGE.-Montpellier, France.

REMARKS.-Said to be in same group with T. globosa, T. oriens, and
T. seminota ( Bergounioux 1938a).



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 207

Testudo ( Testudo) fshensiensis Wiman

Testudo shensiensis Wiman 1930, p. 28, pl. 5, figs. 1-16.
Testudo shansiensis Wiman 1930, p. 36 ( not a typographical error).

TypE.-Institute of Vertebrate Paleontology and Palebanthropology
( Peking ); a shell.

TYPE LOCALITY AND HORIZON.-Fu-Ku-Hsein, Shensi Province, China;
Pontian faunal age, Early Pliocene.

GEOLOGIC RANGE.-Early Pliocene.
GEOGRAPHIC RANGE.-Shensi and Shansi Provinces, China.
REMARKS.-In antiqua-graeca phyletic line according to Glaessner

( 1933), which see for discussion of T. shansiensis and T. shensiensis.
Yeh (1963a) and Gilmore ( 1931) describe additional specimens.

Testudo ( Testudo) f sphaerica Wiman

Testudo sphaerica Wiman 1930, p. 33, figs. 2-26.
Terrapene sinica Yourig 1950, p. 2, figs. 1-2 ( Pliocene, China).
Testudo sinica Mlynarski 1955a, p. 164 ( Pliocene, China).
Testudo Yushensis Yeh 1963a, p. 40 ( Pliocene, China).
Testudo honanensis Yeh 1963a, p. 40, pl. XIII ( Pliocene, China).
Geochelone shaerica Mlynarski 1968, p. 95 ( typographical error).

TypE.-Institute of Vertebrate Paleontology and Paleoanthropology
( Peking ); a shell.

TypE LoCALITY AND HORIZON.-Pao-Te-Chou, Shansi Province, China;
Early Pliocene.

GEOLOGIC RANGE.-Early Pliocene.
GEOGRAPHIC RANGE.-Shansi and Kansu Provinces, North China.
REMARKS.-Not a typical member of the antiqua-graeca line, accord-

ing to Mlynarski ( 1955a); perhaps closely related to T. ( Agrionemys)
hors#eldi. Probably conspecific with some other described Pliocene spe-
cies from North China. Mlynarski ( 1968) places it in the genus Geo-
chelone, but its characters seem more like those of Testudo, where I pre-
fer to keep it. The type of T. gushensis Yeh is placed here in view of its
similarity in morphology and geologic age.

Testudo ( Testudo) tauttoensis Szalai

Testudo suttoensis Szalai 1932, p. 222 ( manuscript name).
Testudo suttoensis Szalai 1934, p. 131.
Testudo graeca Kormos 1932, p. 3 ( part).

TypE.-National Museum of Hungary; humerus, femur, coracoid
and parts of plastron.

TypE LoCALI'rY AND HORIZON.-SUttO, Komarom County, Hungary;
Middle Pleistocene.



208 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

GEOLOGIC RANGE,-Middle Pleistocene.
GEOGRAPHIC RANGE.-Hungary.
REMARKS.-A poorly defned form close to T. marginata ( Szalai

1934) and in the -graeca-group" ( Szalai 1936).

Testudo ( Testudo) f szalai Mlynarski
Testudo szalaii Mlynarski 1955b, p. 164.

TypE.-Polish Zoologic Institute ( Krakow); part of the posterior por-
tion of the shell.

- TYPE LOCALITY AND HORIZON.-Bone breccia of Weze, near Dzial-
oszyn, Lodz Region, Poland; Late Astian and/or Early Villafranchian
faunal ages, Late Pliocene and/or Early Pleistocene (perhaps a mixed
fauna).

GEOLOGIC RANGE.-Late Pliocene or Early Pleistocene.
GEOGRAPHIC RANGE,-Poland.
REMARKS.-Close to T. hermanni according to Mlynarski ( 1955b).

In the antiqua-graeca phyletic line and perhaps a synonym of T. globosa
according to Mlynarski ( 1962).

Testudo ( Testudo ) ttunhuanensis Yeh
Testudinidae Bohlin 1953, p. 63.
Testudo tunhuanensis Yeh 1963a, p. 42, fig. 23-24, pl. XIV, 1-2.

TypE.-Institute of Vertebrate Paleontology and Paleoanthropology
( Peking); two cotypes: a complete plastron and partial carapace.

TYPE LOCALITY AND HORIZON.-Taben-buluk, Tunhuang, Kansu,
China; Miocene?

GEOLOGIC RANGE.-Probably Miocene in age.
GEOGRAPHIC RANGE.-Northern China.
REMARKS.-Probably close to T. sphaerica.

Testudo ( Testudo) tturgaica Riabinin
Testudo turgaica Riabinin 1926, p. 54, fig. 2 , pl. 4 .

TypE.-Geological Museum, Academy of Sciences ( Leningrad ); a
partial shell.

TypE LoCALITY AND HoRIZON.-Dzhilanchik River, near the Kushuka
Wintering Station ( Kugaly-Dzhar Land Section) Turgai area, U.S.S.R.;
Chattian faunal age, Late Oligocene.

GEOLOGIC RANGE.-Late Oligocene.
GEOGRAPHIC RANGE.-Turgai area, U.S.S.R.
REMARKS.-In antiqua-graeca phyletic line according to Glaessner

(1933).



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 209

Genus Uncertain

?Testudo fcastrensis Bergounioux
Testudo castrensis Bergounioux 1935, p. 53.

TypE.-Museum d'Histoire Naturelle de Toulouse; a carapace.
TypE LOCALITY AND HORIZON.-Near Castres, Tam Department,

France; Ludian faunal agd, Late Eocene.
GEOLOGIC RANGE.-Late Eocene.
GEOGRAPHIC RANGE.-France.
REMARKS.-Close to Geochelone richardi according to Bergounioux

( 1938a). If correct, this species is probably a Geochelone.

?Testudo tchienfutungensis Yeh
Testudinidae sp. Bohlin 1953, p. 99, pl. IX, figs. 3-4.
Testudo chienfutungensis Yeh 1963a, p, 100, figs. 25-26, pl. XV, 1, 2.

TYPE.-Institute of Vertebrate Paleontology and Paleoanthropology
( Peking ); a nearly complete carapace.

TypE LOCALITY AND HORIzoN.-Taben-buluk, Tunhuang, Kansu,
China; ?Miocene.

GEOLOGIC RANGE.-?Miocene.
GEOGRAPHIC RANGE.-Type locality only.
REMARKS.-The very narrow shell and presence of intergular scutes

suggest that this turtle may not even be a land tortoise. Both Bohlin
( 1953) and Yeh ( 1963a) consider these additional scutes as abnormalities.

?Testudo fcomptoni ( Bell)
Emys comptoni Bell In Owen 1849.
Homopus comptoni Lydekker 1889a, p. 91.
Testudo comptoni Loveridge and Williams 1957, pp. 218-353.

TypE.-British Museum ( Natural History ); a partial shell.
TypE LOCALITY AND HoRIzoN.-Isle of Sheppey, England; London

Clay, Cuisian faunal age, Early Eocene.
GEOLOGIC RANGE.-Early Eocene.
GEOGRAPHIC RANGE.-Isle of Sheppey, England.
REMARKS.-Fora discussion of relationships see Loveridge and Wil-

liams ( 1957), who consider it close to T. scutella.

?Testudo tpt!/chogastroides Reinach
Testudo ptychogastroides Reinach 1900, p . 19 .

TypE.-?Frankfurt Museum, Germany; shell fragment.



210 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

TYPE LOCALITY AND HORIZON.-Near Frankfurt a. M., Germany;
Burdigalian faunal age, Early Miocene.

GEOLOGIC RANGE.-Early Miocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-Probably not a member of the genus Testudo.

?Testudo treidlii Hoernes

Testudo feidlii Hoernes 1892, p. 243.
"Testudo" C Ocadia) reidlii Siebenrock 1915, p. 360 [sic].

TypE.-Vienna Museum; internal cast of shell.
TYPE LoCALITY AND HORIZON.-Oligocene ( Sotzka layer) in Trifail.
GEOLOGIC RANGE.-Oligocene.
GEOGRAPHIC RANGE.-Austria.

REMARKS.-Redescribed by Teppner ( 1913) who placed it in the

antiqua-graeca line. Possibly an emydid rather than a true tortoise, ac-
cording to Siebenrock ( 1915) and Riabinin ( 1926), with which I concur.

?Testudo tscutella ( Meyer)

Emys scutella Meyer 1845, p. 17, pl. 7, fig. 2.
Homopus scutella Lydekker 1889a, p. 91.
Testudo scutella Loveridge and Williams 1957, p. 353.

TypE.-Teyler Museum ( Haarlem ); a partial carapace.
TypE LOCALITY AND HORIzoN.-Switzerland; (?) Late Miocene.
GEOLOGIC RANGE.-( ?) Late Miocene.
GEOGRAPHIC RANGE.-Switzerland.
REMARKS.-Thought by Lydekker ( 1889a ) to be close to Homopus

areolatus. For discussion of relationships see Loveridge and Williams
( 1957), who considered it close to Testudo comptoni.

?Testudo istehlini Reinach

Testudo stehlini Reinach 1900, p . 113 .

TypE.-Naturhistorischen Museum ( Basel, Switzerland); natural
cast of most of carapace, some carapacial bones, and most of the plas-
tron.

TypE LOCALITY AND HORIZON.-Mathod, near Yverdon, Vaud Canton,
Switzerland; Stampian faunal age, Middle Oligocene.

GEOLOGIC RANGE.-Middle Oligocene.
GEOGRAPHIC RANGE.-Switzerland.

REMARKS.-See Bram ( 1951) for best description; if his statement is

correct that stehlini is closely related to pgrenaica, then it probably be-
longs in the genus Geochelone.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 211

?Testudo tungia Yeh
Testudo tungia Yeh 1963b, p. 224, Eg. 1, pl. 1.

TypE.-Institute of Vertebrate Paleontology and P.aleoanthropology
( Peking ); a carapace.

TypE LOCALITY AND HORIZON.-Gigantopithecus Cave, Liucheng,
Kwangsi, China; Lower Pleistocene.

GEOLOGIC RANGE.-Pieistocene.
GEOGRAPHIC RANGE.-Kwangsi, China.
REMARKS.-Probably not even a testudinid; perhaps referable to the

genus Cuora.

Genera Inquirendae

Genus fCheirogaster Bergounioux
CheiTogaster Bergounioux 1935, p . 78 .

GENOTYPE.-Cheirogaster maurini Bergounioux ( by monotypy).
DEFINmON.-If correctly described, this is a monotypic genus char-

acterized by lack of entoplastron and contact between gular and pectoral
scutes.

TypE LoCALITY AND HoRIZON.-Gironde, France; Eocene.
GEOLOGIC RANGE.-Eocene.
GEOGRAPHIC RANGE.-Type locality,
REMARKS.-If correctly interpreted the type shell represents a distinct

genus. The presumed distinctive characters are so unusual that a re-
examinati6n of the type material is suggested. Bergounioux ( 1958) cor-
rectly points out that his earlier reconstruction of the type species ( 1935)
is greatly in error when compared with the specimen available.

Cheirogaster tmaurini Bergounioux
Cheirogaster maurini Bergounioux 1935, p. 78.

TypE.-M. Maurin Borbereaux collection ( Gironde, France); a com-
plete shell.

TYPE LOCALITY AND HORIZON.-Gironde, France; Ludian faunal age,
Late Eocene.

GEOLOGIC RANGE.-Late Eocene.
GEOGRAPHIC RANGE.-Type locality.

Cheirogaster farrahonensis Bergounioux
Cheirogaster arrahonensis Bergounioux 1957, p. 1237 C Preliminary notice).
Cheirogaster arrahonensis Bergounioux 1958, p. 175, Egs. 15, 16, pls. 32,33.



212 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

TypE.-Museo de Sabadell ( Spain ); a partial shell.
TypE LOCALrrY AND HORIZON.-Sabadell, Penedes District, Spain;

Vallesian faunal age, Late Miocene.
GEOLOGIC RANGE.-Late Miocene.
GEOGRAPHIC RANGE.-Northeastern Spain.
REMARKS.-Very poorly defined.

Genus t Floridemys Williams

Bystra Hay 1916a, p. 53 ( preoccupied by Bystra Cameron 1902).
Floridemys Williams 19504 p. 27 ( substitute name).

GENOTYPE.-Bystra nanus Hay ( = Floridemys nana [Hay] [by mon-
otypy]).

DEFINmON.-A monotypic genus distinguished by its small size and
a transverse gulo-humeral sulcus.

GEOLOGIC RANGE.-Miocene ( originally thought to be Pliocene).
GEOGRAPHIC RANGE.-Central Florida.
REMARKS.-The Miocene age here assigned to the species is based

on both the stratigraphy at the type locality and additional material from
a Miocene site in Alachua County, Florida.

Floridemys fnana ( Hay)
Bystra nanus Hay 1916a, p. 53, pl. 1.
Floridemys nanus Williams 1950a, p. 27.

TypE-U. S. National Museum; a shell.
TYPE LOCALITY AND HORIZON.-Holder Phosphate Mine, near Inver-

ness, Citrus County, Florida, U.S.A.; Hawthorne Formation, Miocene
( Early?).

GEOLOGIC RANGE.-Miocene ( Early?).
GEOGRAPHIC RANGE.-Central Florida, USA.
REMARKS.-Perhaps congeneric with Stylemys, but the small adult

size ( 105 mm. carapacial length) and the transverse gulo-humeral sulcus
suggests that nana should be retained in a distinct genus. Final judg-
ment will have to await additional material. The ending of the species
name has been changed to correspond with the feminine gender of
Floridemys.

Genus f Kansuchelys Yeh

Kansuchelys Yeh 1963a, p. 28.

GENoTypE.-Kansuchelys chiagukuanensis Yeh.
DEFINITION.-Small to medium tortoises with hexagonal neurals

throughout the series, and a doubly notched epiplastral projection.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 213

GEOLOGIC RANGE.-Tertiary ( Oligocene or Eocene?).
GEOGRAPHIC RANGE.-China.
REMARKS.-A very poorly defined genus, undoubtedly very close to,

and perhaps a synonym of the subgenus Manouria ( Genus Geochelone ).
Almost the entire reference to this genus by Mlynarski ( 1968 ) is incorrect.

Kansuchelys f chiayukuanensis Yeh
Kansuchelys chiat,ukuanensis Yeh 1963a, p. 28.

TYPE.-Institute of Vertebrate Paleontology and Paleoanthropology
( Peking ); a nearly complete shell.

TypE LoCALITY AND HORIZON.-Shih-erh-ma-cheng, north of Hui-hui-
p'u, Chia-yu-kuan, Kansu, China; Tertiary, exact horizon unknown.

GEOLOGIC RANGE.-Oligocene or Eocene?
GEOGRAPHIC RANGE.-Type locality only.
REMARKS.-A good specimen of fossil tortoise and important zoo-

geographically; unfortunately with very poor data.

Kansuchelys t oualis Yeh
Kansuchelys oualis Yeh 1963a, p. 33.

TypE.-Institute of Vertebrate Paleontology and Paleoanthropology
( Peking).

TYPE LOCALITY AND HORIZON.-Unknown, probably from Yushe,
Shansi, China.

GEOLOGIC; RANGE.-Unknown.
GEOGRAPHIC RANGE.-Unknown.
REMARKS.-It is unfortunate that this poorly defined species was ever

described.

Genus f Sinohadrianus Ping
Sinohadrianus Ping 1929, p. 232.

GENOTYPE.-Sinohadrianus sichuanensis Ping.
DEFINmON.-An extinct Asiatic genus with neural plates compara-

tively narrow, most neurals hexagonal; plastron extensively united to
carapace; entoplastron wholly in front of pectoral scutes.

GEOLOGIC RANGE.-Eocene.
GEOGRAPHIC RANGE.-Honan, China and Japan.
REMARKS.-The single imperfect shell of the type displays almost

no distinctive features. If it is a tortoise, it is probably more primitive
than Hadrianus. Yeh ( 1963a) states that it is not close to Hadrianus,
and I agree.



214 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

Sinohadrianus leozensis Sbikama

Sinohadrianus eozensis Shikama 1953, p. 20, figs. 1-4, pl. 2.

TypE.-?Natural Science Museum ( Tokyo); carapace.
TYPE LoCALITY AND HORIZON.-Utosinai Coal Mine, Utosinai, Japan;

Eocene.
GEOLOGIC RANGE.-Eocene.
GEOGRAPHIC RANGE.-Japan.

REMARKS.-Not well defined.

Sinohadrianus tsichuan€nsis Ping

Sinohadrianus sichuanensis Ping 1929, p. 232, figs. 1-2, pls. 1-2.

TypE.-Institute of Paleontology and Paleoanthropology ( Peking);
a poorly preserved shell.

TypE LOCALITY AND HORIZON.-Fan Chuang, Si Chuan, Hsien, Ho-
nan, China; Late Eocene.

GEOLOGIC RANGE.-Late Eocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-See Sinohadrianus.

LITERATURE CITED

Agassiz, L. 1857.-Contributions to the Natural History of the United States, 1(2),
North American Testudinata. pp. 235-452, pls. 1-37.

Ahl, E, 1926.-Ueber eine ausgestorbene Riesenschildkrbte der Insel Teneriffa.
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Almera, D. J. and A. P. Bofill. 1903.-Consideraciones sobra los restos fosiles cua-
ternarios de la caverna de Gracia ( Barcelona). Mem. Real. Acad. Ciences y
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Amprino, R. and G. Godina. 1947.-La struttura delle ossa nei vertebrati... com-
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Andrews, C. W. 1904.-Note on the gigantic land tortoise ( Testudo ammon
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. 1906.-A descriptive catalogue of the Tertiary Vertebrata of the Fayum,
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1914.-On the Lower Miocene Vertebrates from British East Africa
Coll. by Dr. Felix Osward. Quart. Journ. Geoi. Soc., London, 70: 163-186, pls.
27-29.

and H. J. L. Beadnell. 1903.-A preliminary notice of a land tortoise
from the Upper Eocene of the Fayum, Egypt. Geol. Surv., Dept. Public Works
Ministry, Cairo, pp. 1-11, pl. 1.

Annandale, N. 1906.-Testudo baluchiomm, a new species. Journ. Proc. Asiatic
Soc. Bengal, 2( 3): 75-76.

Arambourg, C. and J. Piveteau. 1929.-Les vertebres du pontieu de Salmonique.
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Archer, W. H. 1961.-Tortoises with dual gular shield. African Wild Life, 13( 1):
82.

Auffenberg, W. 1961.-A correction regarding the phalangeal formula of the turtle
Stytemus nebrascensis Leidy. Copeia, 1961 (4): 496-498.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 215

1962a.-The status of Testudo anyangensis Ping. Herpetologica,
18( 1): 58-59.

1962b.-A redescription of Testudo hexagonata Cope. Herpetologica,
18( 1): 25-34.

1962c.-A new species of Geochelone from the Pleistocene of Texas.
Copeia 1962, (3): 627-636.

.. 1962d.-Testudo amphithorax Cope referred to Stylemys. Amer. Mus.
Nov., (2120): 1-10.

1963.-The fossil testudinine turtles of Florida, genera Geochelone
and Floridemys. Bull. Florida State Mus.,7(2): 53-97, fgs. 1-33.

. 1964a.-A new fossil tortoise from the Texas Miocene, with remarks
on the probable geologic history of tortoises in eastern United States. The
Pearse-Sellards Series, Texas Mem. Mus., Univ. Texas, 3: 2-10.

. 1964b.-A first record of breeding colour changes in a tortoise. Misc.
Note, Journ. Bombay Nat. Hist. Soc., 61 (1): 191-192.

. 1964d.-A redefinition of the fossil tortoise genus Stylemys Leidy.
Journ. Paleont., 38: 316-324, Egs. 1-6, pl. 51.

1966a.-A new species of Pliocene tortoise, genus Geochelone from
Florida. Journ. Paleont., 40( 4): 877-882.

1966b.-The carpus of land tortoises ( Testudininae). Bull. Florida
State Mus., 10( 5): 159-191.

1967.-Notes on West Indian tortoises. Herp., 23(1): 34-44, 2 fgs.
1969.-Tortoise behavior and survival. Rand MeNally Patterns of

Life Series, 37 pp.
1971.-A new fossil tortoise, with remarks on the origin of South

American tortoises. Copeia 1971, (1): 106-117.
In press.-The genus Gophems. I  Osteology and relationships of

the recent species. Bull. Florida State Mus.
and W. W. Milstead. 1965.-Reptiles in the Quaternary of North Amer-

ica. pp. 557-568 In Wright and Frey, eds., The Quatefnary of the United States,
A review volume for the VII Congress of the International Association for
Quaternafy Research. Princeton Univ. Press.

and W. G. Weaver. 1969.-Gophems bertandien in southeastern Texas.
Bull. Florida State Mus., 13(3): 146-191.

Axelrod, D. I. 1967.-Quaternary extinctions of large mammals. Univ. California
Publ. Geol. Sci., 74: 1-42.

Bachmayer, F. 1967.-Eine Riesenschildkr6te aus den Altpliozanen schichten von
Pikermi ( Griechenland). Proc. Acad. Athenae, 42: 302-317.

Bannikov, A. G. 1947.-( Ontogenesis of some adaptive pecularities in the form of
the shell in tortoises). C. R. Acad, Sci,, Moscow, 58 (4): 709-712, figs. 1-3.

Bartram, W. 1791.-Travels through North and South Carolina, Georgia, East and
West Florida ... containing an account of the soil and natural productions of
those regions. Philadelphia, 522 pp., 8 pls., 1 map.

Bataller, J. R. 1926.-Estudio de restos fosilis de tortuga recientemente encontrados
en Catalaunica. Bol. Inst. Geol. Espana, 46: 145-162, pls. 1-8.

Bate, D. 1914.-Remains of a gigantic land tortoise ( T. gumnesicus n. sp.) from
the Pleistocene of Menorca. Ceol. Mag ( n.s. ), 4( 1): 100-107, figs. 1-2.

Baur, G. 1889.-The gigantic land tortoises of the Galapagos Islands. Amen Nat.,
23: 1039-1057.

1891.-The pelvis of the Testudinata, with notes on the evolution of
the pelvis in general. Journ. Morph., 4: 345-359, figs. 1-13.

Bazhanov, V. S. and N. A. Pigulevsky. 1955.-( On some peculiarities of the age
of the Tertiary tortoise, Testudo kegenika Khozatsky). Acad. Nauk U.S.S.R.,
Mat. Istorri Fauna. Flor. Kazakhstan, 1: 87-94, fgs. 1-2, pl. 1.

Beck, R. H. 1903.-In the home of the giant tortoise. 7th Ann. Rept., New York
Zoological Society, 7: 1-17.



216 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

Beer, G. R. de. 1926.-Studies on the vertebrate head. II. The orbitotemporal re-
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1974 AUFFENBERC: FOSSIL TORTOISE CHECKLIST 217

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1974 AUFFENBERG. FOSSIL TORTOISE CHECKLIST 219

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220 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

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1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 221

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222 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

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Soc. London, Vol. 165: 251-284, pls. 33-45.

1877a.-The gigantic land tortoises (living and extinct) in the col-
lection of the British Museum. 96 pp., pls. 1-54.

1877b.-Description of the living and extinct races of gigantic land-
tortoises. Pts. III & IV. Abstract. Proc. Zool. Soc. London, 25: 506-507.

1896.-Testudo ephippium. Nov. Zool., 3(4): 329-334, pls. XX-XXII.
Haberlandt, G. 1876.-Oeber Testudo praeceps n. sp., die erste fossile Landschild-

kr6te des Wiener Beckens. Jarhb. k. k. Geol. Reichsamst., 26: 243-248, pl. 16.
Hacker, G. and C. Schumacher. 1955.-Die muskeln und faszieren des Mundbodens

bei Testudo graeca. Anat. Anz., 21(4): 52-58.
Haddon, A. C. 1879.-On the extinct land-tortoises of Mauritius and Rodriquez.

Trans. Linn. Soc. London, Ser. 2,2: 155-163, pl. 13.
Hay, Oliver Perry. 1899a.-Descriptions of two new species of tortoises from the

Tertiary of the United States. Proc. U.S. Nat. Mus., 22: 21-24, pls. 4-6.
1899b.-On the nomenclature of certain American fossil vertebrates.

Amer. Geol., 24: 345-349.
1902a.-Bibliography and catalogue of fossil vertebrates of North

America. Bull. U.S. Geol. Sum, 179: 1-449.
. 1902b.-Descriptions of two new species of tortoises from the Tertiary

of the United States. Proc. U.S. Nat. Mus., 22( 1181) : 21-24, pls. 4-6.
1902c.-Descriptions of two species of extinct tortoises, one new.

Proc. Acad. Nat. Sci. Phila.: 383-388.
1903.-Description of a new genus and species of tortoise from the

Jurassic of Colorado. Ann. Carnegie Mus., 2: 201-204.
19042.-On some fossil turtles belonging to the Marsh Collection in

Yale University Museum. Amer. Journ. Sci., 4( 18): 261-275, pls. 11-16, Eg. 5.
1904b.-A new gigantic tortoise from the Miocene of Colorado. Sci-

ence. 19: 503-504.
1907.-Descriptions of new turtles of the genus Testudo, collected

from the Miocene by the Carnegie Museum; together with a description of the
skull of St!/lemvs. Ann. Carnegie Mus., ( 1906) 4: 15-20, figs. 1-11.

1908.-The fossil turtles of North America. Publ., Carnegie Inst.
Wash., (75): 1-555, Egs. 1-704, pls. 1-113.

1916a.-Descriptions of some Floridian fossil vertebrates, belonging
mostly to the Pleistocene. 8th Ann. Rept. Fla. Geol. Surv.: 39-76.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 223

1916b.-Description of some fossil vertebrate found in Texas. Bull.
Univ. of Texas. no. 71: 3-24, pls. 1-4.

1917a.-Vertebrata mostly from stratum no. 3 at Vero, Florida; to-
gether with descriptions of new species. 9th Ann. Rept. Florida State Geol.
Surv.: 43-68.

1917b.-On a collection of fossil vertebrates made by Dr. F. W.
Cragin in the Equus beds of Kansas. Kansas Univ. Sci. Bull., 10 (4): 39-51.

1920.-Descriptions of some Pleistocene vertebrates found in the
United States. Proc. U.S. Nat. Mus., 58: 83-146.

1923.-Characteristics of sundry fossil vertebrates. Pan-Amer. Geol.
( Iowa), 39: 114-120.

1924.-The Pleistocene of the middle region of North America and its
vertebrated animals. Publ. Carnegie Inst. Wash,, 322A: 1-385, fgs. 1-5.

1930.-Second bibliography and catalog of the fossil Vertebrata of
North America. Publ. Carnegie Inst. Wash., 390, vols. 1-2: 1-1074.

Haycroft, J. B. 1891.-The development of the carapace of the Chelonia. Trans.
Royal Soc. Edinburgh, 36( 2): 335-342.

Hediger, H. 1935.-Herpetologische Beobachtungen in Marokko. Verh. Naturf.
Ges. Basel, 46: 1-49, figs. 1-2.

Heller, E. 1903.-Papers from the Hopkins Stanford Galapagos Expedition, 1898-
1899. XIV. Reptilia. Proc. Wash. Acad. Sci., 5: 39-98.

Henri-Martin, G. 1946.-Une tortue fossile dans la Vallee de "Fontechevade"
( Charante). Bull. Soc. Prehistorique Francaise, 43: 86-87.

Hermanni J. 1804.-Observationes Zoologicae, quibus novae compluses ... Opus
post human editit F. L. Hammer. Pais & c. p. 332, Argentorali & Parisus.

Hernandez-Pacheco, E. 1917.-Hallazgo de tortugas gigantescas en el mioceno de
Alcala de Henares. Bol. Real. Soc. Espan. Hist. Nat., ( 17): 194-202.

1921.-Las tortugas fosiles de Palencia. Iberica, 8: 328-330.
1926.-Un nuevo yacimiento de vertebrados fosiles del Mioceno de

Madrid. Bol. Real. Soc. Esp. Hist. Nat., pp. 392-395.
Hewitt, J. 1931.-Descriptions of some African tortoises. Ann. Natal Mus., 6:

461-506, figs. 1-5, pls. 36-38.
1933.-On the Cape species and subspecies of the genus Chersinella

Gray. Pt. I. Ann. Natal Mus., 7: 255-293.
1937.-A note on the relationship of the Cape genera of land tortoises.

South African Journ. Sci., 33: 788-796, fgs. 1-5, pl. 10.
Hibbard, C. W. 1944.-A new land tortoise, Testudo riggsi, from the Middle Plio-

cene of Seward County, Kansas. Univ. Kans. Sci. Bull., 30( 1): 71-76.
1960.-An interpretation of Pliocene and Pleistocene climates in North

America, 62nd Annual Report, Mich. Acad. Sci., Arts & Letters. pp. 5-30.
Hibbard, C. W. and E. S. Riggs. 1949.-Upper Pliocene vertebrates from Keefe

Canyon, Meade County, Kansas. Bull. Geol. Soc. Amer., 60: 829-860, Bgs.
1-11, pls. 1-5.

Hoernes, M. 1892.-Neue Schildkr6tenreste aus steierischen TertiUrablagerungen.
pts. 1-2., Verh. GeoI. Reichsanst., 9: 242-246.

Hoffman, C. K. 1890.-I. Schildkr6ten. In Bronns, Klassen und Ordnungen das
Thierreichs Leipzig, vol. 6, pt. 3: 1-442, pls. 1-48.

Hohenacker, H. 1831.-( Russian turtles). Bull. Nat. Moscow, pp. 361-365.
Holman, J. A. 1958.-The Pleistocene herpetofauna of the Saber-Tooth Cave,

Citrus County, Florida. Copeia, 1958 (4): 276-280.
1959.-Amphibians and reptiles from the Pleistocene ( Illinoisan) of

Williston, Florida. Copeia, 1959 (2): 96-102.
1969.-The Pleistocene amphibians and reptiles of Texas. Publ.

Mich. State Univ., Biol. Ser., 4( 5): 163-192.
Honegger, R. E. 1964.-Beobachtungen an der Spaltenschildkr6te in Ost-Afrika.

Nat. Mus,, 94(12): 462-470.



224 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

Hooijer, D. A. 1948.-Pleistocene vertebrates from Celebes. II. Testudo margae
sp. nov. Proc. K. Nedl. Akad. Wet. Amsterdam, 51( 9): 1169-1182.

1954.-The Pleistocene vertebrate fauna of the Celebes. Arch. Niederl.
Zool., 13 ( 1): 89-96.

1963.-Geochelone from the Pleistocene of Curacao, Netherlands
Antilles. Copeia, 1963 (3): 579-580.

1971.-A giant land tortoise, Geoch€lone atlas ( Falconer and Cautley)
from the Pleistocene of Timor. I. Konikl. Nederl. Akad. van Watenschappen,
Amsterdam. Paleontology, pp. 504-517, pl. 1-2 ( Reprinted from Proceedings,
Ser. B, 74(5), 1971).

Huene, F. von. 1944.-Ein Anomodontien-Fund am oberan Amazonas. Neues
Jarhb. Min. Geol. Paleont. Stuttgart, Abt. B, 10: 260-265.

Hummel, K. 1935.-Schildkr6ten aus der mitteleozlinen Braunkohle des Geiseltales.
N6va Acta Leop., ( N.S. ), 2(3): 457-483, figs. 1-3, pls. 1-29.

Hutchison, V. H., A. Vinegar, and R. J. Kosh. 1966.-Critical thermal maxima in
turtles. Herpetologica, 22( 1): 32-41.

Jackson, J. B. 1837.-Anatomical description of the Galapagos Tortoise. Journ.
Boston Soc. Nat. Hist., I: 443-464, pls. x.xi.

Jeude, T. W. van Lidth de. 1893.-On a new species of the genus Testudo. Notes
Leyden Mus., 15: 312-314, pl. 9.

1896.-On Testudo emys Schleg. & Mull., and its amnities. Notes
Leyden Mus., 27: 197-204, pls. 5-6.

Johnston, C. S. 1937.-Osteology of Bysmachelys cangonensis, a new turtle from
the Pliocene of Texas. Joum. Geol., 45: 439-447.

Joleaud, L. 1906.-Description des terrains neogenes de la plaine der Contal. Mem.
Acad. Vaucluse, 6  360-362.

Kercado, T. 1835.-( no title). Act. Soc. Linn. Bordeaux, 30: 35.
Khozatsky, L. I. 1941.-( Ecological and Morphological study of the evolution of

the shell of terrestrial tortoises). Univ. Leningrad, ( 234): 1-6.
1945.-( Remains of a sea-turtle discovered in Oligocene deposits of

the Asal region). C. R. ( Doklady) Acad. Sci. USSR ( new series) 49: 52-54.
1947.-( A terrestrial tortoise from maeotic deposits of Crimea). C. R.

( Doklady) Acad. Sci. USSR, 58: 2059-2062, 1 fig.
1948.-Novye vidy road Testudo Linne ( Testudines-Reptilia) iz

pliocena Ukrainy. Bjul Kom. Izuc cev periode Akad. Nauk USSR, ( 11): 92-96.
. 1955.-( Land tortoises from the Neogene of the northern Tian-Shan).

Mat. Fauna Flor. Kazak., 2: 539-544, pls. 1-4.
1959.-( On the body surface temperature of some amphibians and

reptiles). Vestn. Leningrad Univ., ( 21): 92- 105, 10 figs.
and M. Mlynarski. 1966.-Agrionemys-nouveau genre de tortue

terrestris ( Testudinidae ). Bull. Acad. Polonaise Sci., 14(2): 123-125.
Kilias, R. 1957.-Die funktionelt-anatomische und systematische Bedeutung der

Schluern reduktion bei Schildkr6ten. Mittl. Zool. Mus. Berlin, 33: 307-354, figs.
Koch, A. 1902.-Neuere Beitrage zu den geologisch-palaontologischen Ver-

haltnissen des Beocsiner Cementmergels. Fmdtani K6z16ny, 32: 311-322.
1904.-Beschreibung der im Beocsiner Cementmergel vorkommenden

Schildkr6tenreste. Ann. Hist. Mus. Nat. Hungarica, (2): 56-61, pls. 7-8.
Koch, W. 1934.-Lungenfasse und kreislauf der Schildkrdten. Biol. Gen. Wein,

10( 2): 359-382.
Koenig, H. 1825.-Icones fossiles sectilas. 18. [not seenl.
Koerner, H. E. 1940.-The geology and vertebrate paleontology of the Fort Logan

and Deep River formations of Montana. Amer. Journ. Sci., 238( 12): 837-861,
pls. 1-7.

Kormos, T. 1932.-Paleolit-kultura maradvanyok Siittorol. Barlangvilag. II: 3-4.
Krefft, G. 1949.-Die Schildkr6ten. Braunschweig, 68 pp.
Kuhn, 0. 1937.-Die fossilen Reptilien. Berlin, 121 pp., 92 figs.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 225

. 1964.-Fossilium Catalogus, I: Animalia, pt. 107, Testudines. W. Junk:
Gravengage. 299 pA

Kutznetzov, V. V. 1958.-( A large species of land tortoise from the Miocene of the
Turgai depression). Acad. Nauk USSR Mat. Istorii Fauny FIory Kazakhstan, 2:
60-68, fgs. 1-2 (in Russian).

Lamanon, R. de Paul. 1780.-M6moire sur la nature et la position ossemens trouv65
d Aix en Provence, dans le couer l'un roches. Journ. Physiol. Chim. Hist. Nat.
Arts, 16: 468-475.

Lambe, L. M. 1906.-Descriptions of new species of Testudo and Baena, with
remarks on some Cretaceous forms. Ottawa Nat., 19: 187-196, pls. 1-3.

1913.-Description of a new species of Testudo and of a remarkable
specimen of St!/lemys nebrascensis from the Oligocene of Wyoming. Ottawa
Nat., 27: 57-63.

Lanfranco, G. G. 1955.-Reptiles, amphibians of the Maltese Islands. Malta Year
Book, pp. 198-203.

Lartet, E. 1951.-Notice sur la colline de Sansan. Auch. Annuaire dept. Gers., pp.
31-52.

1883.-Poissons et Reptiles du Lac de Tiberiade et de quelques autres
parties de la Syrie. Arch. Mus. Nat. Hist. Lyon, 3: 180-192.

Lataste, F. 1881.-( Description of a new land tortoise). Le Naturaliste, p. 396.
1886.-Description d'une Tortue nouvelle de Haut Senegal ( Homopus

noguevi). Le Naturaliste, 8( 2): 286-287.
1888.-Description d'une Tortue nouvelle originaire du Haut Senegal

( Cinixys dorri, n. sp. ). Le Naturaliste, 10( 2): 164-166.
Laurent, J. M. 1768.-Specimen medicum exhibens synopsin reptilium emendatem

cum experimentis circa venena et antidota reptilium Austrizcorm. Viennze. Jan.
Thorn. Nob. de Tratlnern.

Lecointre, Georges. 1926.-Recherches geologogiques dans la Masetra marocaine.
Mem. Soc. Sci. Nat. Maroc., 14: 1-158, pls. 1-18.

LeConte, F. 1854.-Description of four new species of Kinostemon, Proc. Acad.
Nat. Sci. Phila., 7: 180-190.

Legler, J. M. 1959.-A new tortoise, genus Copherus, from northcentral Mexico.
Univ. Kansas Publ., Mus. Nat. Hist., 11(5): 335-343, pls. 7-8.

Legler, J. M. and R. C. Webb. 1961.-Remarks on a collection of bolson tortoises,
Gopherus flauomarginatus. Herpetologica, 17( 1): 26-37.

Lehmann, U. 1957.-Eine jung pleistozane Wirbeltierfauna aus Ostafrica, Mitt.
Geol. Staatsinst., 26: 100-140, figs. 1-26, pls. 8-10.

Leidy, J. 185la.-( On a new species of fossil tortoise). Proc. Acad. Nat. Sci.
Phila., 5: 172-173.

185lb.-( Fossil tortoises from Nebrasca Territory). Proc. Acad. Nat.
Sci. Phila., pp. 326-327.

1852a.-( Notes on fossils from the Nebrasca Territory). Proc. Acad.
Nat. Sci. Phila., 6: 34-60.

1852b.-( Fossil tortoises from Nebrasca). 6th Ann. Rept. Smithsonian
Inst. 1851, p. 65.

1858.-Notice of extinct Vertebrata from the valley of the Niobrara
River. Proc. Acad. Nat. Sci. Phila., pp. 20-29.

1868.-Notice of some vertebrate remains from the West Indian Is-
lands. Proc. Acad. Nat. Sci. Phila., pp. 178-180.

187la.-Remarks on a fossil Testudo from Wyoming and on supposed
fossil turtle eggs. Proc. Acad. Nat. Sci. Phila., pp. 154-155.

187lb.-Remarks on fossil vertebrates from Wyoming. Amer. Journ.
Sci., 2(3): 372-373.

1871c.-Remarks on fossils from Oregon. Proc. Acad. Nat. Sci. Phila.,
pp. 247-248.



226 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

1873.-Contributions to the extinct vertebrate fauna of the Western
Territories. U.S. Geol. Sum Terr., 1: 1-358, pls. 1-37.

1877.-Description of vertebrate remains, chiefly from the phosphate
beds of South Carolina. Journ. Acad. Nat. Sci. Phila., 7: 1-258.

1889.-Description of vertebrate remains from Peace Creek, Florida.
Trans. Wagner Free Inst., 2: 13-17.

Leith-Adams, A. 1877.-A gigantic land tortoise and a small freshwater species from
the ossiferous caverns of Malta. Quart. Journ. Geol. Soc. 33: 177-191.

Lichtenstein, M. H. C. 1823.-Verzeichniss der Dubletten des Zoologischen Museums
der ... Universitat ... Berlin. 118 pp.

Lindholm, W. A. 1929.-Revidiertes Verzeichniss der Gattungen der rezenten Schild-
kr6ten nebst Notizen zur Nomenklatur einiger. Arten, Zool. Anz., 81: 275-295.

1931.-Ober eine angeblicke Testudo aus Sudchina. Zool. Anz., 97:
27-30.

Linnaeus, C. 1758.-Systema Naturae. Holmiae, 10th Ed., 1: 1-824.
1766.-Systema Naturae. Halae Magdeburgicae, 12th Ed., 1: 1-532.

Loomis, F. G. 1909.-Turtles from the upper Harrison beds. Amer. Journ. Sci.,
(ser. 4), 28: 17-26, figs. 1-9.

1927.-A giant tortoise from Florida. Amer. Journ. Sci., ( ser. 5),
( 12 ) : 435-439, figs. 1-2.

Loveridge, A. 1935.-Scientific results of an expedition to rain forest regions in
eastern Africa. I. New reptiles and amphibians from East Africa. Bull. Mus.
Comp. Zool., 79: 1-19.

1942.-Scientific results of a fourth expedition to forest areas in East
and Central Africa. IV. Reptiles. Bull. Mus. Comp. Zool., 91: 237-273, pls. 1-6.

Loveridge, A. and E. E. Williams. 1957.-Revision of the African tortoises and
turtles of the Suborder Cryptodira. Bull. Mus. Comp. Zool., 115: 163-557, pls.
1-18.

Lydekker, R. 1880.-A sketch of the history of the fossil Vertebrata of India.
Journ. Asiatic Soc. Bengal, 69 (2) : 8-40.

1889a.-Catalogue of the fossil Reptilia and Amphibia in the British
Museum, Pt. 3. Chelonia. London, pp. 1-239, figs. 1-53.

1889b.-On the land tortoises of the Siwaliks. Records Geol. Surv.
India, 22(4): 209-212, figs. 1-3.

Lynn, W. G. 1937.-Variation in scutes and plates in the box-turtle, Terrapene
ca,olina. Amen Natl., 71: 421-426.

Lynn, W. G. and M. C. Ullrich. 1950.-Experimental production of shell abnormal-
ities in turtles. Copeia, 1950 ( 4): 253-262.

Maack, G. A. 1869.-Die bis ietzt bekannten fossilen Schildkr6ten ... bei Kelheim
( Bayern ) und Hannover. . . . Paleontologr., 18: 193-338, pls. 33-40.

Macarovici, N. 1930.-Sur une Testudo bessarabica Riab. trouvee dans les couches
Meotiques de Cioburciu. Acad. Roumain., Sect. Sci., 12: 1-2.

Macarovici, N. and St. Vancea. 1960.-Sur les restes de Tortues de la fauna de
Malusteni de la Moldavie meridionale. ( R. P. Roumania). Anal. Stiint. Univ.,
"Al. I. Cuza." Din Jasi, n.s., sect. 2, ( 6,)  377-386,4 pls.

Marsh, O. C. 1890.-Notice of some extinct Testudinata. Amer. Journ. Sci., ( Series
3), 40: 177-179, pls. 7-8.

Matthew, W. D. 1924.-Third contribution to the Snake Creek Fauna. Bull. Amer.
Mus. Nat. Hist., 50: 59-210, figs. 1-63.

Matthew, W. D. and W. Granger. 1923.-The fauna of the Ardyn Obo formation.
Amer. Mus. Novitates, ( 98): 1-5.

McDowell, S. B., Jr. 1961.-On the major arterial canals in the ear-region of the
testudinoid turtles and the classification of the Testudinoides. Bull. Mus. Comp.
Zool., 125(2): 23-39.

Medem, F. 1960.-Datos zoo-geograficos y ecologicos sobre los crocodylia y Tes-
tudinata de Ios Rios Amazonas, Putumayo, y Caquita. Caldasi, 8( 38): 341-351.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 227

1962.-La distribucion geografica y ecologica de los Crocodilyia y
Testudinata en el Departamento del Choco. Acad. Colombinao Ciencias Exactas,
11(44): 279-303, figs. 1-56.

Merrem, B. 1820.-Versuch eines Systems der Amphibien. Tent. Syst. Amphibiorum
Marburg, pp. 1-191, pl. 1.

Merriam, J. C. 1919.-Tertiary mammalian fauna of the Mohave Desert. Bull.
Dept. Geol. Univ. California, pp. 450-533.

Mertens, R. 1936.-Eine bemerkenswerte variation des Schildkr6ten panzers. Isis,
1934, 36( 1): 15-19.

1937.-Reptilien und Amphibien aus dem sudlichen Inner-Afrika.
Abhl. Senck. Naturf. Ges., ( 435) : 1-23.

1946.-Ober einige mediterrane Schildkr6ten-Rassen. Senck., 27( 4-6):
111-118, pls. 1-3.

1956.-Ober reptilienbastarde, II. Senck. Biol., 37: 383-394, figs. 1-3.
and Lorenz Muller. 1928.-Liste der Amphibien und Reptilien Europa.

Abh. Senckenberg. Naturf. Ges. Frankfurt a.M., Band 41: 1-53.
and H. Wermuth. 1955.-Die rezenten Schildkr6ten, Krokodile und

Brukenechsen. Zool. Yahrb., Abt. Syst., 83: 323-440.
Meyer, H. von. 1832.-Palaeologica zur Geschichte der Erde und ihre Geschopfe.

pp. 1-560. Frankfurt a.M.
1834.-Die fossilen Z~hne und Knochen und ihre Ablagerungen in der

Gegend von Georgensmund in Bayern. Suppl. Abht. Mus. Senckenb.,(4): 1-121.
1845.-Zur Fauna der Vorwelt. Fossile Saugethiere, Vogel und Rep-

tilien aus dem Molasse-Mergel Oeningen. Frankfurt a.M., pp. 1-231, pls. 1-12.
1858.-( Briefl. Mitteilung). Neues Jb. Mineral 1858: 296-301.

( no title).
1865.-( Briefl. Mitteilung). Neues Jahrb. Mineral 1865: 603-610

( no title).
1867.-Individuelle abweichungen bei Testudo antiqua und Emys

europaea. Paleontogr., ( 15): 201-221, Bgs. 33-35.
Miller, J. F. 1776-1782.-Various subjects of natural history, wherein are delineated

birds, animals, and many curious plants, etc. 60 col. pls., pp. 1-10. Letterpress:
London.

Miller, L. 1932.-Notes on the desert tortoise ( Testudo agassizii). Trans. San
Diego Soc. Nat. Hist., 7( 18) : 187-208, pls. 10-11.

1942.-A Pleistocene tortoise from the McKittrick Asphalt. Trans.
San Diego Soc. Nat. Hist., 9( 38) : 439-442.

1955.-Further observations on the desert tortoise, Gopherus agassizii
of California. Copeia, 1947 (1): 113-118,2 figs.

Milne-Edwards, A. 1868.-( In Grandidier) Sur des d6couvertes zoologiques faites
r6cemment a Madagascar par M . Alfred Grandidier . C . R. Acad . Sci ., ( Paris ),
pp. 1165-1167.

Milstead, W. W. 1956.-Fossil turtles of Friesenhahn Cave, Texas, with the descrip-
tion of a new species of Testudo. Copeia, 1956 (3): 162-171.

Mlynarski, M. 1955a.-Z6wie z. pliocenu Polski. Acta. Geol. Polonica, 5( 2): 161-
214, figs. 1-20.

1955b.-The systematic position of the Pliocene turtle from Tienshui,
Kansu ( N6rth China ). Acta, Paleont. Sinica, 3(3): 159-165.

1956.-Studies on the morphology of the shell of recent and fossil
tortoises. I-II. Acta. Zool. Cracoviensia, 1( 1): 1-14.

1962.-Notes on the amphibian and reptilian fauna of the Polish
Pliocene and Early Pleistocene. Acta. Zool. Cracoviensia, 7( 11): 177-194, pl. 14.

1966a.-Die fossilen Schildkr6ten in den ungarischen Sammlungen.
Acta Zool. Cracoviensia, 11 (8): 224-288, figs. 1-15, pls. 15-21.

. 1966b.-Morphology of the shell of Agrionemys horsfieldii ( Gray 1844 )
( Testudines, Reptilia). Acta. Biol. Cracoviensia Zool., IX: 219-223.



228 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

1968.-Results of the Polish-Mongolian Palaeontological Expeditions,
Part I. Notes on the Tortoises from the Tertiary of Mongolia. Palaeont.
Polonica, ( 19): 85-97.

1969a.-Fossile Schildkr6ten. Neue Brehm-Bucherei. Verlag-Luther-
stadt. pp. 1-128.

1969b.-Remarks on the fossil Chelonians from Malusteni ... Acta.
Zool. Cracoviensia, 14( 7): 151-161.

Mojsisovics, A. V. 1888.-Zoogeographische Notizen uber Sud-Ungarn Zuleich ein
3 Nachtrag Zur "Fauna von Bellye und Darda." Reptiles and Batrachians. In
Zoo Record. pp. 241-247.

Moreno, F. P. 1889.-Breve resefia de los progressos del Museo La Plata, durente
el segundo semestre de 1888. Boll. Mus. La Plata, 2: 1-44.

Mukerjee, P. N. 1949.-A fossil tortoise shell from the Worli Hill, Bombay. Proc.
34th Indian Sci. Congr., (3): 152.

Nafiz, H. and A. Malik. 1933.-Vertebres fossiles de Kucukeekmece. Bull. Fas.
Sci. Istanbule, ( 3-4): 1-120, pls. i-16.

Nikolsky, A. M. 1896.-Diagnoses Reptilium et Amphibiorum novorum in Persia.
Ann. Mus. Zool. Acad. Sci., St. Petersburg, 1: 369-372.

Nopsca, F. 1926a.-Heredity and evolution. Proc. Zool. Soc. London, pp. 633-665,
figs. 1-9.

1926b.-Osteologia reptilium fossilium et recentium. Fossilium Cata-
logus. Pt. 27, Berlin, pp. 1-391.

1931.-Osteologie reptillium fossilium et recentium. Fossilium Cat-
alogus. I. Animalia, Pt. 50. pp. 1-62.

Obst, Fritz Jurgen and Herwart Ambrosius. 1971.-Taxonomische Studien an euro-
paischen Landschildkr6ten ( Reptilia: Testudinidae) mit serologische-immun-
ologischen Methoden. Zool. Abh., Staatliches Mus. fur Tierkunde, Dresden, Bd.
30( 22): 299-331, Egs. 1-11.

(r

and W. Meusel. 1965.-Die landschildkr6ten Europas. Ziemsen:
Wittenberg, pA 1-56, figs. 1-45.

Oelrich, T. M. 1950.-A new Testudo from Madison County, Montana. Contr. Mus.
Paleont. Univ. Michigan, 8(4): 43-58, Egs. 1-4, pl. 1.

1952.-A new Testudo from the Upper Pliocene of Kansas, with addi-
tional notes on associated Rexroad mammals. Trans. Kansas Acad. Sci., 55(3):
300-311.

1957.-The status of the Upper Pliocene turtle, Testudo turgida Cope.
Journ, Paleont., 31: 228-241, figs. 1-6.

Orbigny, Alcide D. d'. 1847.-Voyage dans l'Amerique Meridionale, etc. ... Tome
Cinquime, lere partie: Reptiles. Bertraud and Levrault, Paris and Strasbourg.

Owens, Richard. 1841.-Description of the remains of six specimens of marine
turtles ( Chelones) from the London Clay of Sheppey and Harwich. Proc. Geol.
Soc. London, III: 570-578.

Pallas, P.S. 1811-42.-Zoographia Rosso-Asiatica, sistem omnium animalium in ex-
tenso Imperio Rosso . . . observatorum . . . atque icones plurimorum. ( Animalia
monocaudia scu frigidi sanguinus... reconsente P. S. Pallas. Supplen dis qui-
busdam Ronarum descriptionibus et iconibus imprimis Piscium ( Amtschaticouem
auxit et locupleiut G. T. Filesius ). 3 Tom. illust. St. Petersburg.

Pantanelli, D. 1893.-Testudo amiatae, sp. n. Atti Soc. Toscana Sci. Nat. Pisa, Mem.,
12: 128-138, figs. 1-2 ( 1892).

Parenzan, P. 1932.-Revisione delle specie del gen. Testudo delle Balcania Atti. 1st.
Veneta, 81: 1149-1169.

Parker, G. H. 1901.-Correlated abnormalities in the scutes and boney plates of the
carapace of the sculptured tortoises. Amer. Nat., 35: 17-24.

Parker, W. K. 1868.-A monograph on the structure and development of the shoulder-
girdle and sternum in the Vertebrata. Ray Soc. London, 237 pp.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 229

Parsons, T. S. 1959.-Nasal anatomy and the phylogeny of reptiles. Evolution, 13:
175-157, 4 figs.

Pawley, R. 1968.-The hidden tortoise of Torreon. Turtle Tort. Soc. Journ., Nov.,
20-23, 36-42.

Peters, K. F. 1869-Zur kenntniss Wirbelthiere aus den Miocanschichten von
Eibeswald im Steimark. I, Die Schildkr6tenreste. Denksch. K. K. Akad. Wiss.

(Wien), 29: 1-71. (1868).
Peyer, B. 1942.-Fossile Riesen Schildkr6ten aus der oberen Susswassermolasse der

Umgebung von Zurich, Schweirischen Palaontol. Abh. Sch. Natur. Gesell., pp.
1-42.

Phillips, J. 1871.-Geology of Oxford and the valley of the Thames. Oxford Univ.
Press. 529 pp.

Pictet, F. J. 1845.-Trait6 61ementaire de Paleontologie Francaise. 1st Ed. (Geneva)
2: 1-407, figs. 1-58.

1853.-Trait6 61ementaire de Paleontologie Francaise. 2nd Ed. ( Paris)
1: 1-562, figs. 1-110.

and A. Humbert. 1856.-Monographie des Ch61oniens de la Mollase
Sidsse. Mat. Paleonts. Suiss6, 3( 1) : 1-71, pls. 1-22.·

Ping, C. 1929.-A new Eocene land turtle from Honan. Bull, Geol, Soc, China,
8(3): 231-238, 2 pls., figs. 1-2.

1930.-Notes on the shell of a land tortoise from the ancient ruins of
Annyang. Bull. Fan. Mem. Inst. Biol., 1. 1-4.

Plieninger, T. 1847.-Verzeichnis der Reptilien Wirtembergs. Jahrb. Ver. raten-
land. Natur. 3: 194-208.

Pomel, H. 1846.-Notes sur les animaux fossiles dicouverts dans le d6partment de
l'Allier. Bull. Soc. Geol. France, ser. 2, voI. 4: 378-385, pls. 1-4.

Portis, A. 1879.-Di alcuni fossili terziarii del Peimonte e della Liguria, apparten-
enti all'ordine dei Chelonii. Mem. Reale Accad. Sci. Torino, ( series 2) 32: 113-
134, 1 pl.

1882.-Les ch61oniens de la Mollasse Vaudoise conserv65 dans le
muse6 gdologique de Lausanne. Mem. Soc. Paleont. Suisse, 9: 1-78, pls. 1-29.

1890.-Rettili pliocenici del Val d'Arno Superiore e di alcune altre
localita plioceniche di Tuscana. Florence, pA 1-32, pls. 1-8.

Power, J. H. 1932.-Testudo verreauxii Smith; a study in variation. South African
Journ. Sci., 29: 466-472.

Procter, J. B. 1922.-A study of the remarkable tortoise, Testudo loveridgii Blgr.,
and the morphology of the cheIonian carapace. Proc. Zool. Soc. London, pp.
483-526, figs. 1-21, pls. 1-3.

Pyburn, W. 1956.-A study of fossil Testudo material from Ingleside, Texas. Univ.
Texas, ( unpublished MS).

Ray, C. E. 1957.-A list, bibliography and index of the fossil vertebrates of Florida.
Florida Geol. Surv., Spec. Publ.,(3): 1-175.

1959.-A sesamoid bone in the jaw musculature of Copherus poly-
phemus. Anat. Ann, 107: 85-91, 4 figs.

Rafnesque, C. S. 1832.-Descriptions of two new genera of turtles of North Amer-
ica. Atlantic Journ. and Friend of Knowledge, Philadelphia. 1:64-65.

Reinach, A. 1900.-Schildkr6tenreste im Mainzer Tertiadrbecken und in benach-
barten ungefahr gleichal-trigen Ablagenungen. Abhl. Senck. Naturf. Ges., 28:
3-135, figs. 1-44.

Riabinin, A. N. 1915.-The tortoises from the Meotian deposits of Bessarabia.
Trav. Mus. Geol. Min., Acad. Sci. Russ, PA 1-16, pls. 1-5, figs. 1-2.

. 1926.-Les Testudo turgaica provenant du Miocene moyen de la region
de Turgai. Acad. Sci. USSR. Trav. Mus. Geol. Leningrad, 1: 52-59, fig. 1.

1927.-Un nouveau ch61onien du tertiare inf6rieur des environs du lac
Issyk-Koul. Bull Comm. Geol. Leningrad, 46: 193-199, pls. 7-9, Bgs. 1-2.



230 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

1945-Une tortue du Meotien de la Crime6. Annu. Soc. Paleon. Russe,
12: 126-132, pls. 1-2.

1946.-Recent ftndings of fossil reptiles in the Crimea. Priroda, 11:
65-66, fgs. 1-2.

Richmond, N. D. 1964.-The mechanical functions of the testudinata plastron.
Amer. Midl. Nat. 72( 1): 50-56.

Rodrigues, J. B. 1892.-Les reptiles fossiles de la vall6e de l'Amazone. Vellosia.
2: 41-52.

Roger, O. 1902.-Wirbeltierreste aus dem Obermiocan der bayerische-schwabischen
Hochebene. Pt. 4. Bericht Naturw. Ver. f. Schwaben-Neuburg ( Ausburg), 35:
1-63, pls. 1-3.

Romer, A. S. 1945.-Vertebrate Paleontology. Univ. Chicago Press, Chicago, 687
pp., 377 Egs.

1956.-Osteology of the reptiles. Univ. Chicago Press, Chicago, 772
pp. 248 Bgs.

Rothschild, L. W. 1901.-On a new land tortoise from the Galapagos Islands. Nov.
Zool., 8: 372.

1902.-Description of a new species of gigantic land turtle from the
Galapagos Islands. Nov. Zool., 9: 619.

1903.-Description of a new species of gigantic tortoises from Inde-
fatigable Island. Nov. Zool., 9: 119.

1906.-A new species of giant tortoise. Nov. Zool., 13: 753-754.
. 1915.-Giant tortoises in the Tring Museum. Nov. Zool., 22: 421-437.

Roverto, C. 1914.-Los Estrados Araucanos y sus F6siles. Ann. Mus. Nac. Hist. Nat.
Buenos Aires, 25: 1-249, pls. 1-31.

Royo y Gomez, J. 1935.-Los grandes tortugas del Seudodiluvial Castellano. Bol.
Soc. Esp. Hist. Nat., 35(9): 463-486, figs. 1-6, pls. 47-54.

Ruckes, H. 1929a.-Studies in chelonian osteology. Pt. I. Truss and arch analogies
in chelonian pelves. Ann. New York Acad. Sci., 31: 31-80.

1929b.-Ibid. Pt. II. The morphological relationships between the
girdles, ribs and carapace. Ann. New York Acad. Sci., 31: 81-120.

. 1937.-The lateral arcades of certain emydids and testudinids. Herpe-
tologica. 1(4): 97-103.

Rutimeyer, L. 1874.-Ueber den Bau von Schale and Schadel bei lebenden und
fossilen Schildkr6ten, als Beitrag zu einer palaontologischen Ceschichte dieser
Thiergruppe. Verh, naturforsch. Gesellech. Basel, 6: 1-137.

Sacco, F. 1889.-I. Cheloni astiani del Piemonte. Mem. Accad. Real. Sci. Torino,
39 (2) : 427-461, pls. 1-2.

Sauzier, T. 1893.-Les Tortues de Terre gigantesques des Mascareignes et de cer-
taines autres iles de la mer des Indes. pp. 1-32.

Scalabrini, P. 1884.-Conferencia el 6 de Enero en la Seccion del Parana. Bol.
Inst. Georg. Argentina. 5: 15-19 [no title].

Schepers, G. W. H. 1938.-The external morphology of the brain of Testudo
geometrica. Journ. Anat., 72: 535-555.

1939.-The blood vascular system of the brain of Testudo geometrica.
Journ. Anat., 73: 451-495, figs. 1-8.

1948.-Evolution of the forebrain. The fundamental anatomy of the
telencephalon, with special reference to that of Testudo geometrica. Cape Town,
212 pp., 250 figs.

Schlegel, H. 1841.-Bemerkungen uber die in der Regentschaft Algier gesammelten
Amphibien. pp. 106-139. In Wagner, Reisen in der Regentschaft Algier in den
Jah~en 1836, 1837, 1838. Leipzig, 3: 1-296.

Schlegel, H. and S. Muller. 1840.-Over de Schildpadden van den Indischen Archi-
pel ... pp. 1-138. In Temminck, Verhandelingen ... natuurlijke ges. Neder-
landsche Overzeesche Bezittingen, published in parts from 1838-1844.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 231

Schmidt, K. P. 1953.-A checklist Of the North American amphibians and reptiles.
Univ. Chicago Press, 215 pp,

Schmidt-Nielson, K. and P. J. Bentley. 1966.-Desert tortoise Gopherus agassizii
cutaneous water loss. Sci., 154( 3750): 911.

Schneider, J. G. 1784.-Sammulungen Vermischt. Abhl. Zool. pp. 1-136.
Schoepff, J. D. 1792.-Historia Testudinum. ArIangae, pp. 1-136, pls. 1-32.
Schreiber, A 1875.-Herpetologia Europaea. F. Vieweg und Sohn, Braunschweig.

pp. 545-563. ( Tortoise part).
Schumacher, C. H. 1954.-Beitrage zur Kiefermuskulatur der Schildkr6ten. I

Mitteilung. Univ. Greifswald. Math. Nat., 3 ( 6-7 ): 457-518, 13 pls., 25 figs.
1955.-Ibid, Parts II and III Mitteilung. Univ. Creifswald. Math. -

Nat., 4(5): 501-523, and 4( 6-7 ) : 559-601, 7 pls., 8 figs.
Schweigger, A. F. 1812.-Monographiae Cheloniorum. Konigsb. Arch. Naturw.

Math., 1: 271-368, and 406«458.
Sclater, P. L. 1870.-Remarks on the animals lately described by Dr. Gray as

Testudo chiliensis and Ateles bartletti. Proc. Zool. Soc. London, pp. 470-472.
Segura, P. A. 1944.-Estudio de la primero especie nueva de tortuga fosil de Costa

Rica con algunas generalidades sombre el orden Testudines. Escuela de Far-
macia Guatemala, 6(73-74): 9-29, (75-76): 16-25, (77-78): 13-14 C continued
over several parts).

Sellards, E. H. 1916.-A new tortoise and a supplementary note on the gavial,
Tomistoma americana. Amer. Journ. Sci., 42: 235-240, Egs. 1-3.

Shah, R. V. and V. B. Patel. 1964.-Myology of the chelonian pectoral appendage.
Jour. Animal Morph. and Physiol., 11 (1): 58-84.

Shaler, N. S. 1888.-Habits of the great southern tortoise. Pop. Sci. Monthly, 34:
37-42.

Shaw, C, 1802.-General Zoology, or Systematic Natural History. 3. Amphibia.
London, 614 pp., 140 pls.

Shikama, T. 1953.-On a new land turtle from the Paleogene of Hokkaido. Trans.
Paleont. Soc. Japan, 9: 19-26, pls, 1-2, figs. 1-4,

Siebenrock, F. 1897.-Das Kopfskelet der Schildkr6ten. Sitzb. d. mathein. -Naturw.
Classe, 56 ( 1): 245-327, figs. 1-6.

1898.-Ueber den Bau und die Entwicklung des Zungenbein-Appar-
ates der Schildkr6ten. Ann. K. K. Naturhist. Hofmuseum, 23 (4) : 424-435.

. 1899.-Ober den Kehlkopf und die Luftr6hre der Schildkr6ten. Sitz.
Kaiser Akad. Wiss. Wien, 108( 7): 563-595, pl. 1.

1900.-Der Zungenbeinapparat und Kehlkopf sammt Luftr6hre von
Testudo calcarata Schneid. Sitz. Kais. Akad. Wiss. Wien, 109 (1): 1-8, pl. 1.

1903a-Ober zwei seltene und eine neue Schildkr6te des Berliner
Museums. Anz. Akad. Wiss, Wien, 40: 185-186 ( abstract of 1903b).

1903b.-Ober zwei seltene und eine neue Schildkr6te des Berliner
Museums. Sitz. Akad. Wiss. Wien, 112 (1): 439-445, pl. 1.

1904.-Ober partielle Hemmungs-Erscheinungen bei der Bildung einer
Ruckenshale von Testudo tornieri Siebenr. Sitz. Akad. Wiss. Wien, 113: 29-34.

1909.-Synopsis der rezenten Schildkr6ten mit Berucksichtigung der
in historischer Zeit ausgestorbenen Arten. Zool. Jahrb., Syst. Suppl., 10: 427-
618.

1910.-Schildkr6ten aus Sud-und Sudwestafrika ... Sitz. Akad. Wiss.
Wien, 119 ( 1) : 693-720, pls. 1-4.

1906.-Zur Kenntnis der mediterranen Testudo-Arten und uber ihre
Verbreitung in Europa. Zoolol. Anzeiger., ( 30): 847-854.

1915.-Testudo kalksbergensis Toula aus dem Leitha-Gebirge. Jb.
K. K. Reichsanstalt, 64: 351-361, pl. 18 ( 1914).

Sieglbauer, F. 1909.-Zur Anatomie der Schildkr6ten extremitUt. Arch. f. Anat. u.
Physiol., Anat. Abt.: 183-280.



232 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

Simionesco, I. 1930.-Les vertebres pliocenes de Malusteni-Roumaine. Acad. Roum.
Publ. Fond. V. Adamachi, 9( 49): 1-15.

Simonetta, A. 1960.-Distribuzione e signiScata dell'organo paratimpanico del
Vitali. Atti Soc. Tosc. Sci. Nat. 66B: 39-55.

Simpson, G. G. 1942.-A Miocene tortoise from Patagonia. Amer. Mus. Nov.,
(1209): 1-6, figs. 1-2.

1943.-Turtles and the origin of the fauna of Latin America. Amer.
Journ. Sci., 241( 7): 413-419.

Sinclair, W. J. 1903.-A new tortoise from the auriferous gravels of California.
Bull. Geol. Univ. California,(3): 243-248, figs. 1-2.

Smets, G. 1885.-Notes sur trois Testudininides de l'Afrique Australe. Ann. Soc.
Sci. Bruelles, 10: 1-11.

Smith, M. 1922.-Notes on reptiles and batrachians from Siam and Indo-China ( No.
1). Journ. Nat. Hist. Soc. Siam, 4(4): 203-214, pl. 8.

1930.-Reptilia and Amphibia of the Malay peninsula. Bull. Raffles
Mus., (3): 1-149.

1931.-Fauna of British India. Vol. I. Reptilia and Amphibia, 256 pp.
Spix, J. B. 1824.-Species novae Testudinum ... Brasilium. pp. 1-42.
Staesche, K. 1931.-Die Schildkr6ten des Steinheimer Beckens. A. Testudinidae.

Paleontr., Suppl., 8(2): 1-17, figs, 1-4, pls. 1-5.
1961.-Beobachtungen am Panzer von Testudo graeca and Testudo

hermanni. Stuttgarter Beitrage z. Naturf., (74): 1-16, fgs. 1-27.
Stammer, A. 1959.-Vergleichende Anatomie und Mikroskopische Innervation der

Augenmuskeln von Reptilien. Acta Biol. Szeged., ( N.S.) 5( 1-2): 125-141, 1 pl.,
3 figs.

Stefano, G. de. 1902a.-Stylemys bottii, n. f. Rivist. Ital. Paleont., 8: 72-76, pl. 7.
1902b.-Cheloniani fossili cenozoici. Bolt Soc, Geol. Ital., 21: 263-

304, pls. 9-11.
Stejneger, L. 1893.-Reptiles and batrachians of the Death Valley Expedition.

North American Fauna, 7: 161.
Stoddart, D. R. and C. A. Wright. 1967.-eography and ecology of Aldabra Atoll.

Atoll Research Bull., ( 118 ): 11-53.
Strain, W. S. 1966.-Blancan mammalian fauna and Pleistocene formations, Huds-

peth County, Texas. Bull. Texas Mem. Mus., ( 10): 1-31.
Strauch, A. 1862.-Chelonologische Studien mit besonderer Beziehung auf die

Schildkr6tensammlung der Kaiserlichen Akademie der Wissenschaften zu St.
Petersburg. Mem. Acad. Imper. Sci. St. Petersburg, (7),4(7): 1-196, pl. 1.

Stromer, E. V. 1926.-Reste Land und Susswasser-Bewohnender Wirbeltiere aus
den Diament-feldern Deutsch Sudwestafrikas, Ch. 2. In Kaiser, "Die Diamenten-
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Sukheswala, R. N. 1947.-A fossil tortoise ( Testudo Zeithii) from the inter-trappeans
of the Worli Hill, Bombay. Proc. 33rd Indian Sci. Congress, (3): 97 ( abstract).

Syevertzov. 1873.-Unfortunately, the original descriptions have not been seen.
Szalai, T. 1930.-Bionomische und methodologisch-sy#tematische Untersuchungen

an rezenten und fossilen Testudinaten. Palaeobiol.,(3): 347-364, pls. 21-24,
I fig.

1932.-Verzeichnis der Ungarischen Testudinaten. F6ld. K6zlony,
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1933.-Schildkr6tenstudien. 1. Testudo schaferi nov, sp., eine Pliozan
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1934.-Die fossilen Schildkr6ten Ungarns. Folia Zool. et Hydrobiol.
Riga, 6(2): 97-142, pls. 1-5.

1935.-Testudo lambrechti Szalai, Folia Zool. et Hydrobiol. Riga, 6(2):
175-178.

1936.-Testudo strandi nov. sp., eine Riesenschildkr6ten aus dem
Mioziin von Surdopusp6ki C Ungarn). Festachrift fur Embrik Strand, Riga, 1:78,
figs. 1-2.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 233

1938.-Fossile Testudo-Reste aus dem Pleistozan Maltas. Annales
Musei Nat. Hungarica Min., Geol., Palaeont., 31: 158-164, figs. 1-2.

Tagliaferro, N. 1914.-On the occurrence of a new gigantic land tortoise at Cor-
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Temminck, C. J. 1840.-Verhandelingen ... natuurlijke ges. Nederlandsche Over-
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and H. Schlegel. 1835.-Reptilia. in Siebold, 1833-1838. Leyden, pp.
1-144, pls. 1-8.

Teppner, W. 1913.-Testudo nedli R. H6ernes. Centralbl. Min. Geol., etc. Yahr-
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Thenius, E. 1952.-Die Schildkr6ten ( Testudinata ) aus dem Unterplioziin von
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Theobald, W. 1879.-On a marginal bone 6f an undescribed tortoise from the
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Thomnson, J S. 1932.-The anatomy of the tortoise. Sci. Proc. Royal Dublin Soc.,
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Thunberg, C. P. 1785-88.-Svenska Vetenskapsakademien, Stockholm. Hondlinger.
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1795.-Travels in Europe, Africa, and Asia, made between the years
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Tinkle, D. W. 1959.-Aspects of shell morphology of North American turtles. Yr. Bk.,
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Tornier, G. 1896.-Die Kriechthiere Deutsch-ost-Afrika. Berlin, 164 pp., 11 figs.,
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Toula, F. 1896.-Ueber neue Wirbeltierreste aus Tertiar 6esterreichs und Rumelieus.
Zeitschr. Deut. Geol. Ges., 48: 915-920, figs., 1-2.

Vaillant, L. 1885a.-Sur une Tortue terrestre d'espece nouvelle, rapporte par M.
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1885b.-Remarques complementaires sur les Tortues gigantesques de
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1889.-Dessins inedits de Cheloniens tires des Manscrits de Cammeron.
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1904.-La livre6 ndotdsique de la Tortue sillon6 ( Testudo calcafata
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1905.-Variations observdes sur le crane chez le Testudo radiata
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Vah Denburgh, J. 1914.-The gigantic land tortoises of the Galapagos Archipelago.
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Van Lidth de Jeude, Th . W. 1898.-On abnormal pectoral shields in Testudo
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Venzmer, J. 1918.-Testudo iberia, var. bicaudalis. Ges, Nat. Freunde Berlin, p.
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1920.-Zool. Anz. 51 ( 1-2): 289 ( title unknown).
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Wagler, J. 1833.-Descriptions et Icones Amphibiorum. Pt. 2, pls. 13-14.
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nach naturlichen Urbildern. Leipzig, pp. 1-132.
Walker, W. F. 1947.-The development of the shoulder region of the turtle,



234 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

Chrvsemys picta marginata, with special reference to the primary musculature.
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Walls, G. L. 1942.-The vertebrate eye and its adaptive radiation. Bull. Cran-
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Wark, A. F. 1929.-A new giant tortoise from the Pliocene of Florida. Amer. Journ.
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Webb, S. D. 1969.-Extinction-origination equilibria in late Cenozoic mammals of
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Weiss, C. S. 1830.-Ceber das sudliche Endes des Gebirgszuges von Brazilien in
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Wermuth, H. 1952.-Testudo hermanni robertmertensi n. subsp. und ihre Vorkom-
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1956.-Versuch der Deutung einiger bisher uberschener Schildkr6ten.
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1958.-Status und Nomenklatur der Maurischen Landschildkr6te,
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Werner, F. 1899.-Beitrage zur Kenntnis der Reptilien und Batrachien-Fauna der
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Wettstein-Westersheim, 0. 1924.-Ober Testudo tornieri Siebr. und Testudo love-
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Wieland, G. K, 1900.-Observations on certain well known stages of the evolution
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1-14.

Williams, E. E. 1950a.-Testudo cubensis and the evolution of western hemisphere
tortoises. Bull. Amer. Mus. Nat. Hist., 95( 1) : 1-36, pls. 1-8, Ags. 1-2.

195Ob.-Variation and selection in the cervical central articulations of
living turtles. Bull. Amen Mus. Nat. Hist., 94(9)  505-561, figs. 1-20.

1952.-A new ~ fossil tortoise from Mona Island, West Indies,  and a
tentative arrangement of the tortoises of the world. Bull. Amer. Mus. Nat. Hist.,
99: 541-560, pls. 1-4, figs. 14.

1953a.-Fossils and the distribution of Chelyid turtles. I. Hydraspis
leithii ( Carter) in the Eocene of India is a Pelomedusid. Brevoria, ( 13 ) : 1-8.

1953b.-A new fossil tortoise from the Thomas Farm Miocene of
Florida. Bull. Mus. Comp. Zool., 107: 534-554, pls. 1-4, figs. 1-4.

1959.-Cervical ribs in turtles. Brevoria ( 101) : 1-12.
1960.-Two species of tortoises in northern South America. Breviora,

(120): 1-12.
Williston, S. W. 1898.-Miocene Edentates. Science, 2(8) : 132.

1925.-The osteology of the reptiles. Cambridge, pp. 1:300.
Wilson, J. W. 1950.-Cope's types of fossil reptiles in the collection of the Bureau

of Economic Geology, the University of Texas. Journ. Paleont., 24: 113-115, figs.
1-3.

Wiman, C. 1930.-Fossile Schildkr6ten aus China. Paleon. Sinica ( Peking), Series
C, 6(.3): 28-33, figs. 1-16.

Woodbury, A. M. and R. Hardy. 1948.-Studies of the desert tortoise, Gopherus
agassizii. Ecol. Monogr., 18( 2): 145-200, pls.

Wu, H. W. 1943.-Notes on the plastron of Testudo emus from the ruins of Shang
dynasty of Anyang. Sinensia, Nanking, 14( 1-6): 107-109.

Wussow, W. 1916.-Meine Erfchrungen mit Testudo horsfieldi, Wochenschrift
Aquar. Terrer. Kunde, 13: 170.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 235

Yeh, H. 1963a.-Fossil turtles of China. Palaeontologia Sinica, ( 150): 1-122.
1963b.-A new Quaternary Testudo from the Gigantopithecus Cave,

Liucheng, Kwangsi. Vert. Palasiatica, 7(3): 224-229.
Young, C. C. 1950.-Notes on a new turtle from Kanshu, China. Paleont. Novitates,

Acad. Sinica, (8): 1-5, figs. 1-2.
Zangerl, R. 1939.-The homology of the shell elements in turtles. Journ. Morphol.,

65: 383-406.
1957.-A parietal foramin in the skull of a recent turtle, Proc, Zool,

Soc. Calcutta. Mukerjee Mem. vol., pp. 269-272, 1 fig., 1 pl.
and R. G. Johnson. 1957.-The nature of shield abnormalities in the

turtle shell. Fieldiana. Geology, 10( 29): 341-362.
Zimmermann-Rollius, S. 1966-Beitrage zur Schildkr6tenfauna der mitteleozanen

Braunkohle des Geiseltales. Hercynia, 4( 1): 83-104.
Zittel, K. A. 1887-90.-Handbuch der Palaeontologie. I. Abt. Palaeozoologic. III.

Vertebrata. Leipzig, 900 pp., 719 8 gs.
Zug, G. R. 1966.-The penial morphology and the relationships of Cryptodiran

turtles. Occ. Papers Mus. Zool; Univ. Mich., ( 647 ): 1-24.



236 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

APPENDIX A
THE GEOLOGIC AND'GEOGRAPHIC DISTRIBUTION OF VALID

LAND TORTOISE SPECIES KNOWN AS FossILS

1,-THE GENus Gopherus.

Area
Time 1 2 3

Recent agassizii berlandieri polyphemus
flauomarginatus

Pleistocene agassizii huecoensis polyphemus
laticaudatus
flavomarginatus
atascosae
cant/onenals

Pliocene pertenuis SP.

Miocene brattstromi 1]an.ms
dehiscus breuistemus
mohauetus copei

undabunus
edae
emiliae

Oligocene neglectus laticuneus
praeextans



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 237

D

C>

0
0



238 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

APPENDIX A ( Co NTINUED)

2.-THE GENUS Stylemys*

Area
Time 1 2 3 4 5

Recent

Pleistocene

Pliocene

genus
Miocene calaverensis (Floddemys?)

capax SP·
conspecta bottiC ?)

Oligocene amphithorax karakoliensis( ?)
ligonia( ?)
nebrascensis

Eocene uintensis

°(?)= generic designation not definite.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 239

6%

0

0



240 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

APPENDIX A (CONTINUED)
3.-THE GENUS Testudo°

Area
Time 1 2 3 4

Recent graeca gra€ca horsfieldi
hermanni hermanni gra€ca
lunellensis marginata kleinmanni

Pleistocene graeca hermanni
seminota suttoensis tungia( ?)
globosa
oriens

Pliocene malai
antiqua cernot>i
seTTesi kucurganica sinica (?)

marcarouiciti shensiensts
sphaerica

illiberatis hypercostata
honanensis

semenensis marmomm hipparionarum marmorum

Miocene celonica darell)skii
scutella( ?) bosporica tunhuanensis
catalaunica kegenika
ptychogastroides(?)
antiqua

Oligocene stehlini (?) alba
lamononi turgaica
denizoti
pusilla
roguesi
reidli( ? )

Eocene doduni
corroyi
cast,ensis( ?)
comptoniC ?)

0(?)- generic designation not certain. Subgeneric designations have been avoided
in view of the unsettled taxonomic situation in both fossil and living populations
belonging to this genus.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 241

0

0
0 9



242 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

APPENDIX A (CONT[NUED)
4.-FoSSIL SPECIES OF THE GENUS Geochelone, SUBGENUS Manouria°

Area
Time 1 2 3 4 5

Recent impressa
emus

Pleistocene

Pliocene puniabiensis

Miocene

Oligocene

Eocene corsoni eocaenica insolitus
gilmorei obailiensis
maiusculus genus
tumida C Kansuchelys?)
utahensis genus

C Sinohadrianus?)

°Genera Kansuchelys and Sinohadrianus may be synonyms.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 243

6%

4, .doe

6 9



244 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

APPENDIX A (CONTINUED)
5.-FOSSIL SPECIES OF THE GENUS Geochetone ( EXCEPTING SUBGENUS Manouria ) 0

( See next page for figure).

Time l 2 3 4 5

Recent pardatis (Ge) pardalis (Ge) elegans (Ge)

Pleistocene amberiacensis (Ge) pardaSs (Ge) pardatis (Ge) allae (Me) kalganensia (He)
burchardi (Ge) cauttevi (Me)
gymnesica (Ge)
robusta (Ge)

Pliocene perpimana (Ge) gchaferi (?) grandia (Ge)
pvrenaica ( Ge) 810anei (T) bessarabica( ?)

ookarkuhni (?)

Miocene botivari ( Ge) crassa (Ge ) namaquensis ( Ge) turgae(Ge)
picteti (Ge) taraktiensis C ?)
vitodu,ana (Ge)
tarteti (7)

.

Oligocene richardi ( Ge) nana(In)
chaileoti ( 7) 1/unnanensis (?)
phosphoritarum(?) meachetica (Ge)

Eocene ammon (Ge) kaiseni (In)
beadnelli ( Ge) ulanensia ( T)
isis (Ge)

•Subgeneric designations in parentheses behind species names, as follows: Al=Aldabrachetvs,
As=Asterochelva, Ca=Caudochetvs, Ch==Chelonoides. Cm=Cymathoic:,8, Cy-c,Zndraspis. Ge
=Geoehetone, He=Hesperotestudo, In==Indoteatudo, Me=Megalochelys, Mo=Monochelys.
T=subgenus inquirendae. *=species became extinct within historic times.



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 245

6 7 8 9 10 11

Emvancorlica(In) pdtastes®(Cy) ca,bonaria(Ch) carbonaria(Ch) chuenms(Ch)
forateni (In) vosmaer€* (Cy) denticulata (Ch) dephan-
dongata(In) Vniphora (As) topus (Ch)
plat~nota ( Ge) rdiata (As)

gigantea ( Al)
indica• (Cy)
mmei.ei* (Al)
grali, (Cy)

ataa (Me) radiata (As) annae (Ca) cubensis (Ch) elata (Ch) chitensis (Ch)
gigantea (Al) crassiscutata(Ca) Bombrerensia(Ch) sellowi (Ch)
abrupta(Al) francisi (Ca) monensis (Mo)
grandidieri  (Al) equicomes (He)
grayi(Cy) incisa ( He)
gadowi (Me) johnstoni (He)

loilsoni (He)

orthowgia (He) gallardoi(Ch)
riggsi ( He)
turgida ( He)
undata ( He)
hat/i (Ca)
alleni (He)
campester (He)
niobrarensis (He)

Inwitata (He) Aesterna (Ch) unngorum
impenaa (He) (Ch)
arenivaga ( He)
kiettiana ( He)
orthopygia (He)
primaeva (He)
vaga (He)
gilberti (He)
f.rri ( He )
william.81(Ca)
tedwhitei (Ca)
milleri ( Ca )
ducatetti (Ca)

quadrata ( ?) coataricensie C ?)
peragrans (?)
cultrata ( ?)
exornata (He)
brontops(Ca)
thompsoniC ?)

longua Km)
achucherti (Cm)



246 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

6%

M

2

49 t:



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 247

INDEX TO SPECIES AND SUBSPECIESS

A burchardi 155
abrupta 148 burnesii 193, 194
agassizii 180, 188
alba 196, 199 C
atlabiatus 139 calarea 139
aHeni 160 calaverensis 191
amberiacen.sis 154 calcarata 157
amiatae 158, 196, 197 campanulata 204
ammon 154 campester 147, 160, 161, 165
amphithorar 131, 188, 176, 190, 192 canetotiana 197
angulata 141 cannetotiana 197
angusticeps 62, 164, 165 canstadiensis 201
annae 145 canyonensis 181, 186
annuliger 173 capax 191
antiqua 189, 168, 171, 175, 178, 196, carbonaria 148, 150

197, 198, 199, 200, 201, 202, 203, carolinus 186
204, 206, 208, 210 carteri 169
a. antiqua 196 castrensis 159,209
a. noviciensis 196 catalaunica 175, 198, 199
a. praeceps 196 cauthley 174

anyangensis 139 cautlegi 178, 174
arachnoides 189 celonica 198
aralensis 140 cernovi 198
arenivaga 160 chaileoti 175
areolatus 187, 188, 210 challoti 175
argentina 148 chiayukyanensis 212, 218
armata 157 chienfutungensis 209
arrahonensis 211 chilensis 134, 135, 137, 148, 149
atascosae 180 claweri 196
atlas 158, 178, 174 commersoni 152
australis 140 comptoni 209, 210

conspecta 191, 192
B copei 182,183

babcocki 157 Corroyi 198
baluchiorum 194 corsoni 129, 169, 171
beadnelli 154 costaricensis 175
belliana 188 couzieri 189, 188
berlandieri 180 crassa 155
bessarabica 175, 198 crassiscutata 133, 144,145, 146
biguttata 156 craveri 196
bipunctata 157 craverii 196
bolicari 154, 155, 158, 160 crawenii 196
bosporica 197 csakvarensis 197
botti 191 csakvariensis 197
bottii 190, 191 cubensis 148
boulengeri 188 culbertsonii 192
boutonii 152 cultrata 176
brattstromi 124, 181, 185,186 cultratus 176
brevisterna 181 cuvieri 140
brevistemus 181 cyclopygia 165
brontops 145 cyclopygius 165
5 Valid taxa are italicized. Page numbers in bold face type indicate the major species
reference.



248 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

D gigantea 142, 148, 144, 170, 176
darewskii 199 g. got,#ei 148
dehiscus 182 gigas 140, 177
demissa 171 gitberti 164 164 179
denizoti 199 gitmord 124, 170,171
denizottii 199 globosa 199, 200, 206, 208
denticulata 148, 150 gopher 186
depressa 124, 181, 186 gou#ei 148
depressus 124, 181, 185 graeca 171,175,178,198, 195,196,197,
despotti 140 198, 199, 200, 201, 202, 208, 204,
distans 145 205,206,207,208,210
doduni 199 g. bettai 201
ducatelli 146, 147 g. boettgeri 201
dumeriliana 189 g. hercegovinensis 201

g. ibera 200
E g. mauritanica 200

ecaudata'200 g. terrestris 200
edae 182, 184 g. zarudnyi 200
elata 149 graii 151
elaverensis 189 graja 204
elegans 142, 158 grandidiefi 148, 144
elephantina 148 grandis 155
elephantopus 148 graweri 196
elongata 167 grayi 151
emiliae 182, 188 gringorum 149
emydoides 169 guentheri 174
emys 168, 169, 170, 172, 175 gymesica 155
enriquesi 201 gymnesicus 155
eocaenica 170
eocenica 170 H
eozensis 214 hadriana 169
equicomes 161 hadrianus 169
erosa 129, 188 hannonensis 202
escheri 196 hagi 146
eurysternum 139 hemispherica 192
exornata 161 hermanni 196, 197, 198,200,201,208,

205, 206, 208
F h. hermanni 201

falconeri 141, 168, 170 h. robertmertemi 201
farri 162 hesterna 131,150
fejervaryi 139 hexagonata 188
femoralis 188 hexagonatus 183, 184
liavomarginatus 180, 183, 184, 187 hipparionum 201
formosa 140 hollandi 182, 184
forsteni 168 homeana 188
francisi 146 honanends 202, 207
frizaciana 140 horsfieldi 193, 194, 195, 202, 207
fusca 169 horsefieldii 194

houzei 140
G huecoensis 184

gadowi 174 hungarica 140
gallardoi 149 hypercostata 202
gaudini 189
geometrica 194, 200 I
geometricus 189 ibera 200
georgicana 200 iberus 194

-

-



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 249

dliberalis 202 lunellensis iberica 204
immensa 162 luxata 169
impensa 162, 165 Iuxatus 141, 168, 169
impressa 169, 170
incisa 160,162 M
indica 151, 152 macarouitii 204

i. perraultii 151 macroeoccygeana 189
i. vosmaeri 150, 152 maiusculus 171

inepta 151 margae 171, 178
innistata 168 , marginata 196, 200, 204, 208
insolita 171 marginatus 196, 200, 204
insolitus 171 m. whitei 200
inusitata 168 marmoreum 205
irregularis 198 marmorum 158, 205
isis 156 maurini 211

mauritanica 200
J mauritonica 200

iohnstoni 163 media 140
mellingi 196

K meshetica 156
kaiseni 168 microtympanum 151
kalganensis 168, 180 milleri 147, 182
kalksbergensis 154, 196 minute 139, 140, 196

k. steinheimensis 197 mira 140
karakolensis 190, 192 mohavense 185
kegenica 202 mohavensis 185
kleinmanni 195 mohauetus 181, 185, 186
klettiana 164, 165 monensis 174
kucurganica 208 munda 139

L N
'amanoni 203 namaquensis 156
amanonii 208 nana 168, 205, 212
amanonis 203 nanus 168, 212
ambrechti 140 nebrascensis 140, 190, 191,192
arnoni 139 neglectus 185, 186
larteti 176 nemoralis 204
ata 192 neoviciensis 196
aticaudata 184 nerandi 140
kiticaudatus 184, 186 niobrarensis 164
aticunae 185 noviciensis 196
piticuneus 185, 187 nurpurensis 140
aurae 189
eberonensis 157 0
eithi 196 obailiensis 172
eithii 139 obtusa 145
emanensis 140 ocalana 145
eptocnemis 151 octonaria 169
igonia 192 octonarius 169
_igonius 192 oculifer 189
ongus 152, 153 olaweri 196
ouisegresmani 146 oregonensis 140
uberonensis 157 oriens 205, 206
uciae 145, 146 orthopygia 161, 162, 165, 167, 179
unanensis 179 o. angusticeps 164, 179unellensis 203 o. orthopygia 165



250 BULLETIN FLORIDA STATE MUSEUM Vol. 18, No. 3

orthopygius 165 radiata 140, 144, 178
osborniana 129, 160,162, 163, 164, 165, radiata fossilis 140

179 reidla 210
oskarkuhni 176 retusa 151
ovalis 218 rexroadensis 161
oweni 192 richardi 154, 155, 158,209

riggsi 166
'P risgoviensis 140

pansa 181, 186 robertmertensi 201
pansus 185, 186 robusta 124, 159, 170, 171
paranensis 140 robustissima 159
pardalis 141, 142, 158, 156, 157, 158, robustus 124, 170, 171

178 rodericensis 152
B babcocki 157 foqued 206
p. pardatis 157 mgosa 163

peltastes 151, 152 rugosus 145
peragrans 177
perpiniana 157, 158,173 S

p. leberonensis 157 sauzieri 151
p. perpiniana 157 scha#eri 177

perraultii 151 schucherti 153
pertenuis 182, 186 scutella 209, 210
phayrei 169 sellardsi 145, 146
phomhoritarum 177 sellovii 150
picteti 158 sellowi 149,150
plana 139 semenensis 206
ptanicauda 141 seminota 206
plat!/nota 153 serresi 206
pliopedemontana 139 serresii 206
polyphaemus 186 shaerica 207
polgphemus 124, 132, 179, 180, 181, shansiensis 207

182,188, 184, 186 shensiensis 202, 207
p. agassizii 180 sichuanensis 218, 214
p. polyphemus 186

pougeti 139 
signata 187
signatus 157, 188

praecedens 186, 187 sinica 207
praeceps 196, 197 sivalensis 173
praeextans 185, 186, 187 sloanei 178
praegraeca 204 snoviana 165

p. ibera 204 sombrerensis 150
praestans 149 st)haerica 207, 208
primaeua 165 spratti 159
promarginata 189, 196 steinheimensis 197
pseudovindobonensis 139 stehlini 210
ptl/chogastroides 209 stellata 142, 153
puniabiensis 170, 172, 175 strandi 140
pusilla 200,205 striata 196
pygmaea 140 strictlandi 189
pyrenaica 158, 175,210 sulcata 141, 148, 153, 157,158

sumeirei 143, 174
Q suttoensis 207

quadrata 177 syrmiensis 155, 197
quadratus 169, 177 szatai 208

szalaii 200
R

racmecskeensis 140



1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 251

T undabunus 187
tabulata campulata 204 undata 162, 165, 166
tarabliensis 178 utahensis 172
tarakliensis 178
tedwhitei 147,162 V
tentorius 189 vaga 167
terrestris 200 vitodurana 155, 158,159, 160
thompsoni 162, 165, 178 uosmaeri 151, 152
tomieri 188, 189
travanconca 168 W
triserrata 151 whitei 200
tumida 172 williamsi 147
tumidus 172 toilsoni 146,167
tungia 211
tunhuanensis 208 y
turgae 159 uniphora 144
turgaica 208 gunnanensis 179
turgida 161, 168, 164, 166, 167, 180 yushensis 207
turmae 159

Z
U zarudnyi 200

uintensis 198 zohalfa 200
utanensis 179 zolkafa 200
undabuna 187



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