• TABLE OF CONTENTS
HIDE
 Copyright
 Front Cover
 Synopsis
 Main
 Literature cited
 Appendix
 Index to species and subspecie...
 Back Matter
 Back Cover






Title: Checklist of fossil land tortoises (Testudinidae)
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Title: Checklist of fossil land tortoises (Testudinidae)
Physical Description: Book
Creator: Auffenberg, Walter.
Publisher: University of Florida,
Copyright Date: 1974
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Table of Contents
    Copyright
        Copyright
    Front Cover
        Front Cover 1
        Front Cover 2
    Synopsis
        Page 121
    Main
        Page 122
        Page 123
        Page 124
        Page 125
        Page 126
        Page 127
        Page 128
        Page 129
        Page 130
        Page 131
        Page 132
        Page 133
        Page 134
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        Page 136
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        Page 139
        Page 140
        Page 141
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        Page 146
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        Page 164
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        Page 169
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        Page 210
        Page 211
        Page 212
        Page 213
    Literature cited
        Page 214
        Page 215
        Page 216
        Page 217
        Page 218
        Page 219
        Page 220
        Page 221
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        Page 228
        Page 229
        Page 230
        Page 231
        Page 232
        Page 233
        Page 234
        Page 235
    Appendix
        Page 236
        Page 237
        Page 238
        Page 239
        Page 240
        Page 241
        Page 242
        Page 243
        Page 244
        Page 245
        Page 246
    Index to species and subspecies
        Page 247
        Page 248
        Page 249
        Page 250
        Page 251
        Page 252
    Back Matter
        Page 253
    Back Cover
        Page 254
Full Text


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of the
FLORIDA STATE MUSEUM
Biological Sciences

VOLUME 18 1974 NUMBER 3





CHECKLIST OF FOSSIL LAND TORTOISES
(TESTUDINIDAE)


WALTER AUFFENBERG


\


UNIVERSITY OF FLORIDA


GAINESVILLE








Numbers of the BULLETIN OF THE FLORIDA STATE MUSEUM, BIOLOGICAL
SC ENCES, are published at irregular intervals. Volumes contain about 300 pages
and are not necessarily completed in any one calendar year.








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This public document was promulgated at an annual cost of $5,227.75 or
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CHECKLIST OF FOSSIL LAND TORTOISES (TESTUDINIDAE)

WALTER AUFFENBERG1


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: Gopherus depressus and Hadrianus robustus, here renamed Gopherus
brattstromi and Geochelone gilmorei respectively. Many species formerly placed in
Testudo (sensu latu) are reallocated to other genera.


1 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, pp. 121-251.




5"'7 0. ;L


,F1O 3

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


TABLE OF CONTENTS

INTRODUCTION --...-.__ ........__.___ .. .. ____ ___ ___ 122
METHODS AND DEFINITIONS ..-----. ----.__ ......--------. .._ 123
ZOOGEOGRAPHY AND ECOLOGY ..._........__---_-------- -__ -__ -. _______ 124
EVOLUTION, PHYLOGENY, AND TRENDS -------..._ ---__ ____._.... ___.. 127
TAXA INCORRECTLY PLACED IN TESTUDINIDAE _-.-........-.. __.._..____ 139
NOMINA NUDA AND UNIDENTIFIABLE MATERIAL ------_- __~__ ----- 140
SYSTEMATICS _---....----.. _-...--..__............___ 140
Genus Acinixys --.-___.--------......_----__ - ------ 140
Genus Chersina ......-------_-- ------_-_-___._ ...... 141
Genus Geochelone .---------------------------_-....... 141
Subgenera
Aldabrachelys .------------ 142 Hesperotestudo -_ ~- 160
Asterochelys ---------.----. 144 Indotestudo---___-------- 167
fCaudochelys ------ 144 Manouria -_______- 168
Chelonoidis .._---.-- 148 fMegalochelys ____ 173
fCylindraspis ..----- ....---------.- 150 +Monachelys __- __ 174
fCymatholcus .-------..-----------...... 152 Subgenera Inquirendae ----__ 175
Geochelone ..----------- --___ _.... 153
Genus Gopherus .. ------__----- --_ .-.. .. 179
Genus Homopus .._-_-___----__ ---- --. .. ---- 187
Genus Kinixys -------188
Genus Malacochersus .------......_- --.. .. _-_-..... ...... . 188
Genus Psammobates --------..-_..-------- --..-.......--- 189
Genus Pyxis ---------.......-_ ------- ------------ --------.. 189
Genus fStylemys -...- -------___...__.......... --------....- ...-- 190
Genus Testudo ---- ----------------..............------ 193
Subgenera Pseudotestudo ------ 195
Agrionemys --.....--.-----------.-.. 194 Testudo __-------------- 195
Genus Uncertain ---_-. ----.-.. ...-----..-..-----------.. . 209
Genera Inquirendae ---_- ----- -------- --- 211
fCheirogaster .-..------_---- ---...... 211 fKansuchelys----.------ ----- 212
fFloridemys ------------------.--------_ 212 tSinohadrianus --- ~- 213
LITERATURE CITED ---------- -----...--- --- 2....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.








AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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 osteological 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 fossils.
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 localities 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 between the genera,
subgenera, or species of fossil and Recent forms. The number and kinds of differ-








BULLETIN FLORIDA STATE MUSEUM


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 f.
(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 polyphemus), Re-
cent of southeastern United States.
Gopherus jdepressus Brattstrom is here renamed Gopherus tbrattstromi in
honor of its discoverer.
B. Hadrianus frobustus Gilmore (1915), Uinta Eocene, Utah (= Geochelone
[Hadrianus] robustaa.
Testudo robusta Leith-Adams (1877), Pleistocene Malta (=Geochelone
[?Geochelone] robustaa.
Hadrianus frobustus Gilmore is here renamed Geochelone igilmorei 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 reflect 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, Malachochersus,
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


Vol. 18, No. 3








AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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 tor-
toises in the middle and northern latitudes 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
(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.


1974




















. t


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


'-r a"








AUFFENBERG: FOSSIL TORTOISE CHECKLIST


extant elephantine tortoises is represented by gigantic continental rela-
tives. Large carnivores existed in these same continental areas from the
Eocene to the Recent' alongside 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 lines.
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, though only near the culmination of a series of climatic changes
that had already greatly reduced the ranges of the giant species of many
different animals (Webb 1969). Man's repeated visits to those Pacific
and Indian Ocean islands formerly harboring innumerable 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) some 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 Stylemys. 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

'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.








BULLETIN FLORIDA STATE MUSEUM


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, 1950b,
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 turtles1). 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

x 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, Williston 1925,
Beer 1926, Nopsca 1926a, 1931, Ruckes 1929a, 1929b, Bolk et al. 1931-8, Thompson 1932, Vers-
luys 1936, Kuhn 1937, Schepers 1938, 1939, 1948, Zangerl 1939, Walls 1942, Romer 1945, 1956,
Walker 1947, Schumacher 1954, 1955, George and Shah 1955a, 1955b, 1959a, 1959b, Golby and
Gamble 1957, Parsons 1959, Williams 1959, McDowell 1961, Shah and Patel 1964, Zug 1966,
Pawley 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.


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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



B L4 8 4 8 4 I^J^<6



C 6> 6> 6> 4 <6 6 <61<6



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

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

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







BULLETIN FLORIDA STATE MUSEUM


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
posterior 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 (Stylemys, Homo-


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


neurals. A. Stylemys famphithorax, Early Oligocene, Colorado, U.S.A. B. Geochelone
fhesterna, 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


1974







BULLETIN FLORIDA STATE MUSEUM


A





















B


















FIGURE 4.-The primitive pleural rib and peripheral pit articulation of the carapace
of pond turtles (A, Chrysemys 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 polyphemus). The distal end of the pleural ribs are
usually weakly developed, or absent.


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


A


FIGURE 5.-Median longitudinal section of the epiplastron. A. Chrysemys scripta
Pleistocene (Emydidae). B. Stylemys famphithorax, Oligocene (Testudinidae). C.
Geochelone fcrassiscutata, Pleistocene (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-
fied 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


1974








BULLETIN FLORIDA STATE MUSEUM


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


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


U r













C D













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 subulnar elements, and lateral migration of
both subulnar and subradial elements accompanying fusion of the me-
dialia (Fig. 9) (Auffenberg 1966b).







BULLETIN FLORIDA STATE MUSEUM


'Cr K




'W27 4

~j2 ~3


B





U

c:3 CD ,


1 W 4
2 3


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


A Bi C

= r U r ulna.














(Fig. 7).
Finidae 9-Major trends thos 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


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


A












B














FIGURE. 10.-The stapedial notch (arrow) is usually open in emydid turtles (A,
Chrysemys 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.








BULLETIN FLORIDA STATE MUSEUM


The land tortoises undoubtedly arose from primitive emydids, prob-
ably toward the close of the Late Cretaceous. The 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 long, 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 into peripheral pits
11) At least prefrontal and frontal scales present
12) Scales on forelimb numerous, not greatly 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 paired
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 morphology 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


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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

TAXA INCORRECTLY PLACED IN THE TESTUDINIDAE1

Testudo fmunda Hay 1920. Pleistocene of Tennessee, U.S.A. (=Terrapene carolina,
see Auffenberg 1963).
Testudo ipromarginata Reinach 1900. Lower Miocene of Frankfort a.m., Germany.
(=Ptychogaster ifrancofurtanus, in part, and T. antiqua, in part, see Glaessner
1935).
Testudo ifejervaryi Szalai 1930. Aquitanian of Salgotarian, Hungary. (=Ptycho-
gaster ifejervaryi, see Glaessner 1933).
Testudo feurysternum Gervais (ex-Pomel) 1836. Lower Miocene of France.
(=Ptychogaster feurysternum, see Braim 1951).
fArchaeochelys pougeti Bergounioux 1938b. (anything organic?).
tColossoemys macrococcygeana Rodrigues 1892. Miocene of Peru. (=Triassic
ammonodont, see Huene 1944).
Testudo flaurea Forster and Becker 1888. (=Ptychogaster flaurae, see Glaessner
1933).
Testudo istricklandi Phillips 1871. ( =Protochelys tstricklandi, see Lydekker 1889a).
Testudo fcalarea Fritsch 1893. Upper part of the Lower Miocene of Eger, Bohemia,
Germany. (=Ptychogaster fcalarea, see Glaessner 1933).
Testudo fleithii Sukheswala 1947. Eocene of India. (= Hydraspis leithii [Carter]
Mukerjee 1949; [ =Carteremys leithii Williams 1953a]).
"Testudo" Iplana Koenig 1825. (=Puppigerus? crassicostatus Owen 1841).
Testudo fanyangensis Ping 1930. Archeologically associated in South China. (=Pseu-
docadia fanyangensis, see Lindholm 1931 and Auffenberg 1962a).
Testudo fminuta Fraas 1870 (part). Miocene of Steinheim, Germany. (=Clemmys
fsteinheimensis Staesche 1931).
Cinixys fcouzieri Bergounioux 1935. Oligocene of France. (= Ptychogaster
fcouzieri).
Emys fgaudini Pictet and Humbert 1856. (= Kinixys [Ptychogaster] igaudini
Portis 1882 [= Ptychogaster igaudini]).
"Testudo" felaverensis Bravard 1858. (=Clemmys fbravardi).
Testudo flamoni (part) Gervais 1859. (=Clemmys fvidali).
Hadrianus fallabiatus Cope 1872c. (=Achilemys fallabiata Hay 1908 =?Emydidae).
Although Kuhn (1964) lists Testudo pseudovindobonensis as appearing on p. 126
of Szalai (1934), he is actually referring to Trionyx pseudovindobonensis Szalai (p.
134). Kuhn also lists Testudo dumeriliana Lartet (1851), which should read Emys
dumeriliana Lartet. His listing of Testudo pliopedemontana probably falls in the
same category. However I have not seen the paper in which this reference was
supposedly made (Sacco 1889).

'Wermuth (1956) discusses the very broad 18th Century use of the generic term "Testudo" for
nontestudinid turtles. Names involved are Testudo boddaerta (=Trionyx cartilaginosis), Testudo
brevi-caudata (=Terrapene c. carolina), Testudo caouana (=Caretta caretta), Testudo caro-
liniana (=Terrapene carolina), Testudo dorsata (=Geoemyda punctularia), Testudo fimbria
(=Chelus fimbriatus), Testudo flava (=Emys blandingi), Testudo granulosa (=Trionyx punc-
tatus), Testudo marina vulgaris (=Chelonia mydas), Testudo meteagris (=Emys blandingi),
Testudo membranacea (=?Trionyx cartilaginosus), Testudo mydas minor (=Lepidochelys o.
kempi), Testudo nasicornis (=Caretta caretta), Testudo planitia (=Macroclemys temminki),
Testudo punctata (=Emys orbicularis), Testudo rubicunda (=Pelomedusa subrufa), Testudo
rugosa (=Chelonia m. japonica), Testudo semimembranacea (=Trionyx sinensis), Testudo ser-
pentina (=Chelydra serpentina), Testudo striata (=Trionyx cartilaginosus and T. triunguis
part), Testudo verrucosa (=Geoemyda punctularia), Testudo viridi-squamosa (=Lepidochelys o.
kempi).


1974








BULLETIN FLORIDA STATE MUSEUM


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 fminuta Bravard 1844. (Szalai 1934).
Testudo media Bravard 1844. (Staesche 1931).
Testudo faralensis Khozatsky 1945.
(Testudo) thouzei Dollo 1912. Only the species name was used originally and later
assumed to be Testudo by Bergounioux (1933b).
Testudo fnurpurensis Meyer 1865. (Kuhn 1964).
Testudo faustralis Moreno 1889. (Williams 1950a).
Testudo trisgoviensis Fraas 1870. (provisional name).
Testudo iformosa Moreno 1889. (Williams 1950a).
Testudo fparanensis 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 inerandi Gray 1831a.
Testudo fcuvieri Fitzinger 1835, ( = Testudo radiata fossilis Meyer 1832; = Testudo
radiata Pictet 1845).
Machrochelys fmira Meyer 1858, ( = Testudo [Macrochelys] mira Zittel 1889).
Stylemys foregonensis Leidy 1871c. (=Testudo foregonensis Leidy 1873;
?Stylemys inebrascensis Cope 1885).
Testudo ffrizaciana Lartet 1851 (=Stylemys ifrizaciana Auffenberg 1964d) (Pictet
1853).
Testudo fdespotti Szalai 1934. (Kuhn 1964).
Testudo fhungarica Szalai 1934.
Testudo flambrechti Szalai 1934.
Testudo tstrandi Szalai 1936. (Mlynarski 1966a).
Testudo igigas Bravard 1844.
Testudo flemanensis Bravard 1844. (Riabinin 1926).
Testudo tracmecskeensis Szalai 1932 (Ms. name), 1934. (Glaessner 1935).
Testudo ipygmaea Lartet 1851, (=Stylemys Jpygmaea 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 enclosed 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).
Pyxis Boulenger 1889, p. 145 (part).
Acinixys Siebenrock 1903b, p. 244-6, pl. 33-34.


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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 1831a, p. 7, 14 (not of Humphreys).
Goniochersus 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
Chersine Merrem 1820, p. 29 (part).
Testudinites Weiss 1830, p. 293.
Geochelone Fitzinger'1835, p. 111.
Megalochelys Falconer and Cautley 1837, p. 358 (part).
Colossochelys Falconer and Cautley 1844, p. 54.
Geoemyda Cantor 1847, p. 2 (part).
Teleopus Le Conte 1854, p. 187 (T. luxatus).
Manouria Gray 1854, p. 134.
Emys Milne-Edwards In Grandidier (part) 1868, p. 1167.
Scapia Gray 1869, p. 169 (S. falconeri).
Stylemys Cope 1870, p. 124 (part).
Peltastes Gray 1870b, p. 655 (part).
Gopher Gray 1870a, p. 190.
Testudo Leidy 1871a, p. 154 (part).
Centrochelys Gray 1872a, p. 5 (T. sulcata Miller, by monotypy).
Hadrianus Cope 1872a, p. 2 (part).
Stigmochelys Gray 1873a, p. 5 (T. pardalis Bell, by monotypy).
Elephantopus Gray 1873b, p. 724 (part).
Eupachemys Leidy 1877, p. 7 (nomen nudum).
Homopus Boettger 1893, p. 8 (part).


1974







BULLETIN FLORIDA STATE MUSEUM


Pampatestudo Lindholm 1929, p. 285 (part).
Megachersine Hewitt 1931, p. 257 (T. pardalis Bell, by original designation).
Gopherus Williams 1950a, p. 30 (part).
GENOTYPE.-TeStudo stellata 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 Aldabrachelys Loveridge and Williams
Aldabrachelys 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.
DEFINITION.- 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.


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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 igigantea
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) aabrupta (Vaillant)
Testudo abrupta Milne-Edwards In Grandidier 1868, p. 1161 (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 LOCALITY 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 fgouffei (Rothschild)
Testudo gouffei 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 Giinther (1877a, 1877b).

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







BULLETIN FLORIDA STATE MUSEUM


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 five 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) Igrandidieri (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
Asterochelys Gray 1873a, p. 4.
Testudo Vaillant 1885a, p. 440.
Geochelone Loveridge and Williams 1957, p. 342.
TYPE SPECIES.-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 1950b, Vaillant 1905, Siebenrock 1897.

Subgenus fCaudochelys Auffenberg
Caudochelys Auffenberg 1963, p. 69 (as subgenus).
TYPE SPECIES.-Testudo crassiscutata Leidy (= Geochelone [Caudo-
chelys] crassiscutata [Leidy]).
DEFINITION.-Extinct Nearctic subdivision of genus Geochelone with


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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) fannae (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) fbrontops (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 (Leidy)
Eupachemys obtusa Leidy 1877, p. 232.
Eupachemys 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.
Gopherus ocalana Williams 1950a, p. 30.
Geochelone distans Ray 1957, p. 126.


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BULLETIN FLORIDA STATE MUSEUM


Geochelone luciae Ray 1957, p. 126.
Geochelone sellardsi Ray 1957, p. 126.
Geochelone (Caudochelys) crassiscutata Auffenberg 1963, p. 70.
TYPE.-U.S. National Museum; parts of plastron, femur and tibia.
TYPE LOCALITY 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) tducatelli (Collins and Lynn)
Testudo ducatelli Collins and Lynn 1936, p. 166, pls. 3-4.
Testudo ducateli Kuhn 1964, p. 116 (typographical error).
Geochelone ducatelli 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.-Late 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 (Caudochelys) thayi (Sellards)
Testudo hayi Sellards 1916, p. 235, fig. 7, 9.
Testudo louisekressmani Wark 1929, p. 401.
Gopherus hayi Williams 1950a, p. 30.
Geochelone (Caudochelys) hayi Auffenberg 1963, p. 78.
Geochelone louisekressmani Auffenberg 1963, p. 79.


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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.-CIose and perhaps ancestral to the Blancan G. campester.

Geochelone (?Caudochelys) fmilleri (Brattstrom)
Testudo miller 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 Bernardino 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) ftedwhitei (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) fwilliamsi Auffenberg
Geochelone williamsi 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).








BULLETIN FLORIDA STATE MUSEUM


Subgenus Chelonoidis Fitzinger
Testudinites Weiss 1830, p. 293.
Chelonoidis Fitzinger 1856, p. 112.
Gopher Gray 1870a, 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 (1950a). 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, Giinther 1875, 1877a, 1896,
Jeude 1896, Siebenrock 1897, Heller 1903, Ruckes 1937, Williams 1950a, Zangerl
1957, Auffenberg 1966b, 1971. Zoogeography: Bauer 1889, Simpson 1943, Auffen-
berg 1971.

Geochelone (Chelonoidis) chilensis (Gray)
Testudo sulcata Dumeril and Bibron 1835, p. 74 (part).
Testudo sulcata d'Orbigny 1847, p. 6 (not of Gmelin).
Testudo (Gopher) chilensis Gray 1870a, p. 190 (Chile, in error).
Testudo argentina Sclater 1870, p. 471 (substitute name for T. chilensis Gray
1870a).
Geochelone chilensis Williams 1960, p. 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.


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TYPE LOCALITY AND HoRIzoN.-Chapepote Springs, Banas de Ciego
Montero, Las Villas Province (= Santa Clara Province), Cuba; Late
(?) Pleistocene.
GEOLOGIC RANGE.-Late Pleistocene.
GEOGRAPHIC RANGE.-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 LOCALITY 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) fgallardoi (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 LOCALITY 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) fgringorum (Simpson)
Testudo gringorum Simpson 1942, p. 1, figs. 1-2.
TYPE.-American Museum of Natural History; a plastron and most
of a carapace.
TYPE LOCALITY AND HORIZON.-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 (Auffenberg 1971).


1974








BULLETIN FLORIDA STATE MUSEUM


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 AND 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) isellowi (Weiss)
Testudinites sellovii Weiss 1830, p. 293.
Testudo sellovii 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) isombrerensis (Leidy)
Emys sombrerensis Leidy 1868, p. 180.
Testudo sombrerensis Williams 1952, p. 552.
Geochelone sombrerensis Auffenberg 1967, p. 35.
TYPE.-Academy of Natural Sciences (Philadelphia) (Lost ?).
TYPE LOCALITY 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
Cylindraspis Fitzinger 1835, p. 112.
TYPE SPECIEs.-Testudo indica vosmaeri Shaw (= Geochelone [Cyl-
indraspis] vosmaeri [Shaw]).


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DEFINITION.-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 a nuchal
scute, undivided supracaudal, 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) findica (Schneider)
Testudo indica Schneider 1784, p. 355.
Testudo indica perraultii Shaw 1802, p. 25.
Chersine retusa 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 (Cylindraspis) fgrayi (Dumeril and Bibron)
Testudo graii Dumeril and Bibron 1835, p. 155.
Testudo inepta Giinther 1873, p. 397 (Mauritius).
Testudo triserrata Giinther 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 sauzieri Gadow 1894, p. 315 (Mauritius).
TYPE.-Museum of Natural History (Paris); a shell.
TYPE LOCALITY.-Unknown, presumably Mauritius Island, Indian
Ocean (Giinther 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. Giinther (1877a) provides an excellent resume of the
skeleton. The data available show that all 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) ipeltastes (Dumeril and Bibron)
Testudo peltastes Dumeril and Bibron 1835, p. 138.
Testudo vosmaeri Fitzinger 1826, p. 1 (part).


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BULLETIN FLORIDA STATE MUSEUM


TYPE.-Museum of Natural History (Paris); dried specimen, less
head and tail.
TYPE LOCALITY.-Rodriguez Island, Mascarene Group, Indian Ocean.
GEOLOGIC RANGE.-Recent, now extinct.
GEOGRAPHIC RANGE.-Rodriguez Island, Indian Ocean.
REMARKS.-Giinther (1877a) and Boulenger (1890c) considered the
specimens of peltastes to be the young of vosmaeri. 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) fvosmaeri (Shaw)
Testudo indica Schoepff 1792, p. 103. (part).
Testudo indica vosmaeri Shaw 1802, p. 27.
Testudo vosmaeri Fitzinger 1826, p. 44.
Testudo rodericensis Giinther 1873, p. 397 (preliminary note).
Testudo boutonii Giinther 1875, p. 43.
Testudo commersoni 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 tCymatholcus Clark
Cymatholcus Clark 1932, p. 132.
GENOTYPE.-Cymatholcus longus Clark 1932.
TYPE LOCALITY 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 longus Clark 1932, p. 132.


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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 RANGE.-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.
Cymatholcus schucherti Williams 1950a, 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 (1950a) 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
Geochelone 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. pardalis Bell).
TYPE SPECIES.-Testudo elegans Schoepff (=Geochelone [Geoche-
lone] elegans [Schoepff]).
DEFINITION.-An Old World subdivision of the genus Ceochelone 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 1950b, Auffenberg 1966b. Fifteen extinct species are
known.


1974







BULLETIN FLORIDA STATE MUSEUM


Geochelone (Geochelone) famberiacensis (Deperet)
Testudo amberiacensis 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.

Geochelone (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) beadnelli (Andrews)
Testudo beadnelli Andrews 1906, p. 285, fig. 91.
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.-Probably conspecific with G. ammon.

Geochelone (Geochelone) tbolivari (Hernandez-Pacheco)
Testudo sp. Hernandez-Pacheco 1917, p. 197.
Testudo bolivari Herandez-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.


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GEOGRAPHIC RANGE.-Spain.
REMARKS.-It seems doubtful that all the specimens figured 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. vitodurana Biedermann.
The best description is by Royo y Gomez (1935).

Geochelone (Geochelone) fburchardi (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) tcrassa (Andrews)
Testudo crassa Andrews 1914, p. 181.
Testudo crassa Szalai 1938, p. 162.
TYPE.-British Museum (Natural History); pieces of the 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 syrmiensis by Simi-
onesco (1930), but this is certainly incorrect.

Geochelone (?Geochelone) Igymnesica (Bate)
Testudo gymnesicus Bate 1914, p. 102, figs. 1-2.







BULLETIN FLORIDA STATE MUSEUM


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) fisis (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) Imeshetica (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) jnamaquensis (Stromer)
Testudo namaquensis Stromer in Kaiser 1926, p. 139.
TYPE.-Berlin Museum of Zoology; a complete plastron.
TYPE LOCALITY 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 Geochelane pardalis.

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


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Testudo bipunctata Gray 1831a, p. 12 (erroneously attributed to Cuvier MS).
Testudo armata Boil (MS) (listed as synonym by Gray 1931a, p. 4 (nomen nudum).
Testudo pardalis Gray 1831a, p. 12.
Geochelone pardalis 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 pardalis babcocki Loveridge 1935, p. 4.
Testudo pardalis pardalis Loveridge 1935, p. 4.
Geochelone pardalis Hewitt 1937, p. 789.
Testudo pardalis pardalis Mertens 1937, p. 5.
Geochelone pardalis babcocki Loveridge and Williams 1957, p. 235.
Geochelone pardalis pardalis Loveridge and Williams 1957, p. 251.
TYPE.-British Museum (Natural History)?; a preserved adult.
TYPE LOCALITY.-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 fleberonensis (Deperet)
Testudo leberonensis 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 HORIZON.-Mount Leberon, west of Cucuron,
France; Pontian faunal age, Early Pliocene.
GEOLOGIC RANGE.-Early Pliocene only? (see remarks below).
GEOGRAPHIC RANGE.-Type locality.
REMARKs.-Peyer (1942) considered leberonensis very close to, and
perhaps a synonym of Geochelone perpiniana, here treated as a separate
subspecies.

Geochelone (Geochelone) perpiniana tperpiniana (Deperet)
Testudo perpiniana Deperet 1885, p. 214, pl. 4.
TYPE.-Museum of Perpignon (Southern France); slightly damaged
specimen.
TYPE LOCALITY AND HoRIzoN.-Near Perpignon, France; Plaisancian
faunal age, Late Pliocene.


1974







BULLETIN FLORIDA STATE MUSEUM


GEOLOGIC RANGE.-Late Pliocene only? (see remarks below).
GEOGRAPHIC RANGE.-France only? (see remarks below).
REMARKS.-G. perpiniana, G. picteti, G. bolivari, and G. vitodurana
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) fpicteti (Biedermann)
Testudo picteti Biedermann 1863, p. 18, pis. 2, 2a.
TYPE.-Winterthur Museum (Switzerland); an almost complete shell.
TYPE LOCALITY 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) ipyrenaica (Deperet and Donnezan)
Testudo pyrenaica 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) jrichardi (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.


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

Geochelone (?Geochelone) robusta (Leith-Adams)
Testudo robusta Leith-Adams 1877, p. 178, pis. 5-6 (not robustus of Gilmore).
Testudo spratti Leith-Adams 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 Xolca, Corradino Hill, Maghlak, Ghar
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 action
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) fturgae (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) fvitodurana (Biedermann)
Testudo vitodurana 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-


1974







BULLETIN FLORIDA STATE MUSEUM


gested that G. vitodurana may be only subspecifically distinct from G.
bolivari.
Subgenus fHesperotestudo Williams
Hesperotestudo Williams 1950a, p. 25 (as subgenus).
TYPE SPECIEs.-Testudo osborniana Hay (= Geochelone [Hespero-
testudo] osborniana [Hay]).
DEFINITION.-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) falleni Auffenberg
Geochelone alleni Auffenberg 1966a, p. 877.
TYPE.-Florida State Museum; shell of adult female.
TYPE LOCALITY AND HoRIzoN.-McGehee 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) tarenivaga Hay
Testudo arenivaga Hay 1907, p. 16, figs. 6-8.
Geochelone arenivaga Auffenberg 1963, p. 93.
TYPE.-Carnegie Museum; pygal and one peripheral.
TYPE LOCALITY AND HORIZON.-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) jcampester (Hay)
Testudo campester Hay 1903, p. 627 (nomen nudum).


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Testudo campester Hay 1908, p. 455, figs. 610-613.
Copherus campester Williams 1950a, p. 30.
Testudo rexroadensis Oelrich 1952, p. 301 (Rexroad fm., Kansas).
Geochelone rexroadensis 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) fequicomes (Hay)
Testudo equicomes Hay 1917b, p. 41, pis. 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 Formation, Rancholabrean faunal age, Sangamon
Interglacial, Pleistocene.
GEOLOGIC RANGE.-( Sangamon) Late Pleistocene.
GEOGRAPHIC RANGE.-Kansas, U.S.A.
REMARKS.-CIose to Geochelone (Hesperotestudo) turgida (see
Auffenberg 1962c, 1963).

Geochelone (?Hesperotestudo) fexornata (Lambe)
Testudo exornata Lambe 1906, p. 187, pl. 3, figs. 1-3.
Geochelone exornata Auffenberg 1962c, p. 635.
TYPE.-Geological Survey of Canada; fragmentary pleural bones.
TYPE LOCALITY 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.


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Geochelone (Hesperotestudo) ifarri (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) Igilberti (Hay)
Xerobates undata Gilbert 1898, p. 143, figs. 1-4.
Testudo gilbert Hay 1899b, p. 349.
Geochelone gilbert 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.
Geochelone (Hesperotestudo) incisa Auffenberg 1963, p. 82.
TYPE.-U. S. National Museum; a xiphiplastron.


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TYPE LOCALITY AND HoRIzoN.-Ocala Lime Company Quarry, near
Ocala, Marion County, Florida, U.S.A.; (?) Sangamon Interglacial, Pleis-
tocene.
GEOLOGIC RANGE.-Pleistocene.
GEOGRAPHIC RANGE.-Florida, U.S.A.
REMARKS.-In Geochelone (Hesperotestudo) turgida evolutionary
line (Auffenberg 1962c, 1963).

Geochelone (Hesperotestudo) \inusitata (Hay)
Testudo inusitata Hay 1907, p. 18.
Testudo innistata Szalai 1930, p. 354 (typographical error).
Gopherus inusitata Williams 1950a, 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 RANGE.-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) fjohnstoni Auffenberg
Testudo rugosa Johnston p. 47 (manuscript name).
Geochelone johnstoni 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) fkalganensis (Gilmore)
Testudo kalganensis Gilmore 1931, p. 247, pl. 9.
Gopherus kalganensis Williams 1950a, p. 22-23.
Geochelone kalganensis 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.


1974







BULLETIN FLORIDA STATE MUSEUM


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

Geochelone (Hesperotestudo) tklettiana (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) \niobrarensis (Leidy)
Testudo (Stylemys) niobrarensis Leidy 1858, p. 29.
Stylemys niobrarensis 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 RANGE.-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 fangusticeps (Matthew)
Testudo angusticeps Matthew 1924, p. 207 (in error).
Testudo orthopygia 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.


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REMARKS.-Perhaps close to the Geochelone osborniana evolution-
arv 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 cyclopygius Cope 1878, p. 394 ("Miocene," Loup Fork, Kansas).
Caryoderma snoviana 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 snoviana Hay 1902a, p. 451.
Geochelone orthopygia Williams 1950a, 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.-Close to Geochelone (Hesperotestudo) campester. In
the G. thompsoni-G. angusticeps-G. osborniana-G. gilbert line, ac-
cording to Matthew (1924). For best description of skeletons see Hay
(1908).

Geochelone (Hesperotestudo) osborniana (Hay)
Testudo osborniana Hay 1904b, p. 503.
Geochelone osborniana Williams 1950a, p. 30.
TYPE.-American Museum of Natural History; a complete shell,
skull, and post cranial skeleton.
TYPE LOCALITY 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. thompsoni-G. angusticeps-G. orthopygia-G. gilbert
evolutionary line (Matthew 1924).

Geochelone (Hesperotestudo) fprimaeva (Oelrich)
Testudo primaeva Oelrich 1950, p. 44.
TYPE.-Univ. of Michigan Museum of Paleontology; shell and a few
girdle and limb elements.








BULLETIN FLORIDA STATE MUSEUM


TYPE LOCALITY AND HORIZON.-Locality 8, west side Sweetwater
Creek, about 2 miles southwest Belmont Park Ranch House, N.E. 1/4
Sec. 32, T8S, R5W, Madison County, Montana, U.S.A.; Early Miocene.
GEOLOGIC RANGE.-Early Miocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKs.-Relationship not clear.

Geochelone (Hesperotestudo) triggsi (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 LOCALITY 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) fturgida (Cope)
Emys turgida Cope 1870, p. 125.
Testudo turgida Cope 1892a, p. 127.
Gopherus turgida Williams 1950a, p. 30.
Geochelone turgida 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.


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TYPE LOCALITY AND HomzoN.-Santa Fe Basin, near Santa Fe,
New Mexico, U.S.A.; Santa Fe series, probably Clarendonian faunal age,
Early Pliocene.
GEOLOGIC RANGE.-Early Pliocene.
GEOGRAPHIC RANGE.-Central New Mexico, U.S.A.
REMARKS.-Closely related to Geochelone orthopygia.

Geochelone (Hesperotestudo) tvaga (Hay)
Testudo vaga Hay 1908, p. 414, pl. 19.
Gopherus vaga Williams 1950a, p. 30.
Geochelone vaga Auffenberg 1964a, p. 6.
TYPE.-American Museum of Natural History; apparently parts of
three individuals.
TYPE LOCALITY 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) \wilsoni (Milstead)
Testudo wilsoni Milstead 1956, p. 168.
Geochelone wilsoni Auffenberg 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 to Early Recent.
GEOGRAPHIC RANGE.-Eastern Oklahoma and Central Texas to New
Mexico, U.S.A.
REMARKs.-Last known member of the important Geochelone tur-
gida evolutionary line (Auffenberg 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]).
DEFINITION.-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.








BULLETIN FLORIDA STATE MUSEUM


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) fkaiseni (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 LOCALITY 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 it has no close relatives.

Geochelone (?Indotestudo) inana (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 HORIZON.-East Mesa, Twin Oboes, Shara
Murun Region, Inner Mongolia; ?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 Miiller 1840, p. 37 (part).
Geoemyda Cantor 1847, p. 2 (part).
Manouria Gray 1852, p. 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 1871a, p. 154 (part).
Hadrianus Cope 1872a, p. 2.
TYPE SPECIES.-Testudo emys Schlegel and Miller (= Geochelone
[Manouria] emys [Schlegel and Miiller]).


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DEFINITION.-A rather primitive Holarctic and Oriental subgenus
of the genus Geochelone, characterized by a long narrow skull with
many emydid 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 1871a, p. 154.
Emys carter Leidy 1871b, 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 quadratus 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 Miller)
Testudo emys Schlegel and Miiller 1840, In Temminck, p. 34 (Sumatra) (see RE-
MARKS).
Geoemyda spinosa Cantor 1847, p. 2 (part).
Testudo emydoides Dumeril and Dumeril 1851 (substitute name for T. emys Schlegel
and Miiller).
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 Giinther 1864, p. 10.







BULLETIN FLORIDA STATE MUSEUM


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 LOCALITY.-Sumatra.
GEOLOGIC RANGE.-Pliocene of the Siwaliks (Lydekker 1889b) to
the Recent.
GEOGRAPHIC 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 (Smith 1922, 1930). Lydekker
(1889b) suggested Geochelone puniabiensis may be close to G. emys.
REFERENCES.-Gray 1855, 1870c, Boulenger 1889, Jeude 1896, Wil-
liams 1950b, Auffenberg 1966b.

Geochelone (Manouria) feocaenica (Hummel)
Testudo eocaenica Hummel 1935, p. 463, pls. 2, 3.
Testudo eocenica Kuhn 1964, p. 117 (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) jgilmorei (new substitute name)
Hadrianus robustus Gilmore 1915, p. 146, pl. 25 (preoccupied by Testudo robusta
Leith-Adams [1877], here placed in Geochelone [?Geochelone]).
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.


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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) finsolitus (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 LOCALITY AND HORIZON.-Promontory Bluff, Mongolian Repub-
lic; 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) Jmargae (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)
to be related to some of the Upper Siwalik species, though this is un-
certain.

Geochelone (Manouria) tmajuscula (Hay)
(?) Hadrianus corsoni Cope 1874, p. 36 (part).
Hadrianus majusculus Hay 1904a, p. 271, pl. 15.
TYPE.-Yale Peabody Museum; a shell.
TYPE LOCALITY AND HonRzoN.-Near Gallina, Rio Arriba County,
New Mexico, U.S.A.; Wasatchian faunal age, Early Eocene.
GEOLOGIC RANGE.-Early Eocene.


1974







BULLETIN FLORIDA STATE MUSEUM


GEOGRAPHIC RANGE.-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, fig. 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) fpunjabiensis (Lydekker)
Testudo punjabiensis Lydekker 1889b, p. 87.
TYPE.-Indian Museum; right and left epiplastra.
TYPE LOCALITY AND HORIZON.-Siwalik Hills, Punjab, India; Plio-
cene (?).
GEOLOGIC RANGE.-Pliocene (?).
GEOGRAPHIC RANGE.-Punjab, India.
REMARKS.-Lydekker (1889b) suggests it may be close to G. emys.

Geochelone (Manouria) ftumida (Hay)
Hadrianus tumidus Hay 1908, p. 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 RANGE.-Utah, U.S.A.
REMARKS.-The ending of the species name has been changed to
correspond with the feminine gender Geochelone.

Geochelone (Manouria) tutahensis (Gilmore)
Hadrianus utahensis Gilmore 1915, p. 148.
TYPE.-Carnegie Museum; plastron and bridge.
TYPE LOCALITY AND HORIZON.-South of Kennedy's Hole, Uinta


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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 fMegalochelys Falconer and Cautley
Megalochelys Falconer and Cautley 1837, p. 358.
Colossochelys Falconer and Cautley 1844, p. 54,
Testudo Lydekker 1880, p. 20 (part).
TYPE SPECIES.-Colossochelys atlas Falconer and Cautley (= Geo-
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) atlas (Falconer and Cautley)
Megalochelys sivalensis Falconer and Cautley 1837, p. 358 (subsequently withdrawn).
Colossochelys atlas Falconer and Cautley 1844, p. 54.
Colossochelys (Megalochelys) 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 margae Hooijer 1948, p. 1169, fig. 1 (Pleistocene, Celebes).
Testudo margae Hooijer 1954, p. 486.
TYPE.-British Museum (Natural History); an epiplastron.
TYPE LOCALITY AND HORIZON.-Siwalik 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.
perpiniana. 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) cautleyi (Lydekker)
Cautlega annuliger Theobald 1879, p. 186 (nomen nudum).


1974







BULLETIN FLORIDA STATE MUSEUM


Testudo cautleyi 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) Igadowi (Van Denburgh)
Testudo guentheri Gadow 1894, p. 320 (invalid name, not of Baur 1889, proposed
for T. sumeirei; 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 fMonachelys Williams
Monachelys Williams 1952, p. 547 (as subgenus).
TYPE SPECIES.-Testudo (Monachelys) 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 first 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) fmonensis (Williams)
Testudo (Monachelys) monensis Williams 1952, p. 547, pls. 44-47.
TYPE.-American Museum of Natural History; skull, parts of shell,
and partial skeleton.


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


TYPE LOCALITY AND HORIZON.-Lirio Cave, Mona Island, West In-
dies; Late Pleistocene.
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 tbessarabica (Riabinin)
Testudo bessarabica Riabinin 1915, p. 6, pls. 1, 3, 4.
TYPE.-Geology Museum, Academy of Sciences (Leningrad); partial
shell.
TYPE LOCALITY 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 punjabiensis and G. emys. Glaessner (1933)
states that it is not in the antiqua-graeca phyletic line. Mlynarski (1969b)
correctly places it in Geochelone.

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

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


1974







BULLETIN FLORIDA STATE MUSEUM


TYPE.-National Museum of Costa Rica; an almost complete shell.
TYPE LOCALITY AND 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 jcultrata (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 LOCALITY AND HORIZON.-Head of Horse Tail Creek, Weld
County, Colorado, U.S.A.; White River Series, Chadronian faunal age,
Early Oligocene.
GEOLOGIC RANGE.-Early Oligocene.
GEOGRAPHIC RANGE.-Northeastern Colorado, U.S.A.

?Geochelone flarteti 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 RANGE.-Late Miocene.
GEOGRAPHIC RANGE.-Sansan, France.
REMARKS.-A giant tortoise, probably referable to the genus Geo-
chelone.

?Geochelone foskarkuhni Mlynarski
Geochelone 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


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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

?Geochelone fperagrans (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.-(?) Oligocene.
GEOGRAPHIC RANGE.-Type locality.
REMARKS.-The relationships of this species are unknown.

?Geochelone jphosphoritarum (Bergounioux)
Testudo phosphoritarum 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 fschafferi (Szalai)
Testudo schafferi Szalai 1933, p. 154 (not Ptychogaster schafferi Glaessner).


1974







BULLETIN FLORIDA STATE MUSEUM


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 isloanei (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.-Closely related to Geochelone radiata or G. pardalis ac-
cording to Lydekker (1889a) though certain of its features suggest it
may be close to Testudo.

?Geochelone ftarakliensis 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.-Affinities 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 tthompsoni (Hay)
Testudo thompsoni Hay 1908, p. 400, pi. 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, U.S.A.; Lower Oreodon beds, Orellan faunal age, Middle Oligo-
cene.
GEOLOGIC RANGE.-Middle Oligocene.
GEOGRAPHIC RANGE.-Western South Dakota.


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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. gilbert, and G. orthopygia, according to Mat-
thew (1924).

?Geochelone fulanensis (Gilmore)
Testudo ulanensis Gilmore 1931, p. 245.
Geochelone ulanensis Mlynarski 1968, p. 96.
Hadrianus ulanensis Chkikvadze 1970b, 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.-Affinities 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 Iyunnanensis (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 Rafinesque
Testudo Bartram 1791, p. 18.
Gopherus Rafinesque 1832, p. 64.
Xerobates Agassiz 1857, p. 446.
Bysmachelys Johnston 1937, p. 439.
GENOTYPE.-Testudo polyphemus Daudin ( = Gopherus polyphemus
[Daudin]).
DEFINITION.-Nearctic genus of tortoises with short cervical verte-







BULLETIN FLORIDA STATE MUSEUM


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 (1950a). 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. flavomarginatus, and G. polyphemus.
REFERENCES.-Zoogeography. Blair 1958, Brattstrom 1961. Mor-
phology. Skeleton: Williams 1950b, Auffenberg 1964b, 1966b, in press;
Skull: Williams 1950a, 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 fatascosae (Hay)
Testudo atascosae Hay 1902c, p. 383.
Gopherus atascosae Hay 1924, p. 247.
TYPE.-Academy of Natural Sciences (Philadelphia); parts of a
plastron.


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TYPE LOCALITY 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 conspecific with it.

Gopherus tbrattstromi (new substitute name)
Gopherus depressus Brattstrom 1961, p. 548 (Preoccupied by Testudo depressa
Guerin-Meneville [1829], here placed in Gopherus 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. pansa, according to Bratt-
strom (1961).

Gopherus fbrevisternus (Loomis)
Testudo brevisterna Loomis 1909, p. 21.
Gopherus brevisterna 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 Icanyonensis (Johnston)
Bysmachelys canyonensis Johnston 1937, p. 440.
Gopherus canyonensis Williams 1950a, p. 21.
TYPE.-Panhandle Historic Plains Museum (Canyon, Texas); a plas-
tron and skeleton (plastron now lost).
TYPE LOCALITY 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.


1974








BULLETIN FLORIDA STATE MUSEUM


GEOGRAPHIC RANGE.-Northwestern Texas and beds of similar age
as far west as southern Arizona, U.S.A.
REMARKS.-Close to C. pertenuis. Close to G. polyphemus accord-
ing to Williams (1952).

Gopherus tcopei (Koerner)
Testudo copei Koerner 1940, p. 838.
Gopherus copei Williams 1950a, p. 30.
TYPE.-Yale Peabody Museum; a shell.
TYPE LOCALITY AND HORIZON.-Sec. 14, R10N, T5E, 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
Gopherus 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
miller (Brattstrom).

Gopherus fedae (Hay)
Testudo edae Hay 1907, p. 19.
Gopherus edae Williams 1950a, 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. hollandi. The best description is by
Hay (1908).


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1974 AUFFENBERG: FOSSIL TORTOISE CHECKLIST 183
Gopherus \emiliae (Hay)
Testudo emiliae Hay 1908, p. 419.
Gopherus emiliae Williams 1950a, p. 30.
TYPE.-American Museum of Natural History; an almost entire shell.
TYPE LOCALITY 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 flavomarginatus Legler
Gopherus polyphemus Duges 1888, p. 146-147 (part).
Gopherus flavomarginata Legler 1959, p. 337.
Gopherus polyphemus flavomarginatus 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 (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 fhexagonatus (Cope)
Testudo hexagonata Cope 1893, p. 77.
Gopherus hexagonata Williams 1950a, 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-








BULLETIN FLORIDA STATE MUSEUM


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

Gopherus fhollandi (Hay)
Testudo hollandi Hay 1907, p. 18.
Gopherus hollandi Williams 1950a, p. 30.
TYPE.-Carnegie Museum; a complete shell.
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. edae.

Gopherus thuecoensis Strain
Gopherus 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 LOCALITY 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 polyphemus 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 laticaudata 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.


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Gopherus flaticuneus (Cope)
Testudo laticunea Cope 1873a, p. 6.
Gopherus laticunea Williams 1950a, p. 30.
TYPE.-American Museum of Natural History; an almost complete
carapace, pelvis, and some limb elements.
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.
GEOLOGIC RANGE.-Early and Middle Oligocene.
GEOGRAPHIC RANGE.-Northeastern Colorado, southeastern Wyoming,
southwestern South Dakota, and western Nebraska, U.S.A.
REMARKS.-Gopherus praeextans and G. neglectus may be synonyms
of G. laticuneus. 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 Gopherus.

Gopherus \mohavetus (Merriam)
Testudo mohavense Merriam 1919, p. 456.
Testudo mohavensis Bataller 1926, p. 155.
Gopherus mohavense Williams 1950a, p. 30.
Gopherus mohavetus Des Lauriers 1965, p. 1.
TYPE.-Univ. of California Museum of Paleontology; a complete
shell.
TYPE LOCALITY AND HORIZON.-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 fneglectus (Brattstrom)
Gopherus neglectus Brattstrom 1961, p. 544, figs. 1-5.
TYPE.-California Institute of Technology; an almost entire shell.
TYPE LOCALITY AND HORIZON.-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.-CIose to G. praeextans and G. laticuneus according to







BULLETIN FLORIDA STATE MUSEUM


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

Gopherus fpansus (Hay)
Testudo pansa Hay 1908, p. 420, pl. 71, figs. 1-2, text figs. 550-1.
Gopherus pansa Williams 1950a, 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. mohavetus,
according to Brattstrom (1961). The ending of the species name has
been changed to correspond with the masculine gender of Gopherus.

Gopherus 1pertenuis (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 polyphemus (Daudin)
Testudo polyphaemus Bartram 1791, p. 18 (nomen nudum).
Testudo polyphemus 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 polyphemus Stejneger 1893, p. 161.
Gopherus carolinus Shaler 1888, p. 37.
Gopherus praecedens Hay 1916a, p. 55 (Late Pleistocene, Vero, Florida).
Gopherus polyphemus polyphemus 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.


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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. flavomarginatus (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 Pleistocene and Recent in age (Weigel 1962).

Gopherus fpraeextans (Lambe)
Testudo praeextans Lambe 1913, p. 61.
Gopherus praeextans Williams 1950a, 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 fundabunus (Loomis)
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








BULLETIN FLORIDA STATE MUSEUM


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 1950b.

Genus Kinixys Bell
Kinixys Bell 1827, p. 398.
Cinothorax Fitzinger 1835, pp. 108, 111, 121.
Cinixys 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).
DEFINITION.-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 170 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 1950b, Kilias 1957, Villiers 1958, Auffenberg
1966b.

Genus Malachochersus Lindholm
Cinicys Tornier 1896, p. 2 (part) [sic] (not of Gray).
Cinixys Procter 1922, p. 515.
Malacochersus Lindholm 1929, p. 285.
Testudo Siebenrock 1903a, p. 185.
GENOTYPE.-Testudo tornieri Siebenrock (= Malacochersus tornieri
[Siebenrock]).
DEFINITION.-An African genus with a persistently fenestrated, very


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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 1950b, 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]).
DEFINITION.-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.
GEOGRAPHIC 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
1950b, Auffenberg 1966b.

Genus Pyxis Bell
Pyxis Bell 1827, p. 395.
?Testudo Boulenger 1889, p. 45.
Bellemys Williams 1950b, p. 553 (in error).
GENOTYPE.-Pyxis arachnoides Bell.
DEFINITION.-A monospecific genus of small tortoise from the Mala-







BULLETIN FLORIDA STATE MUSEUM


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 1873c, Williams 1950b.

Genus tStylemys Leidy
Stylemys Leidy 1951a, p. 173 (not of Maack 1869).
Emys Leidy 1851a, p. 173.
Testudo Leidy 1851b, p. 173.
Geochelone Auffenberg 1963, p. 87 (part).
Gopherus 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.
Stylemys 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.


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GEOLOGIC RANGE.-Early Oligocene.
GEOGRAPHIC RANGE.-Northeastern Colorado and (?) South Dakota,
U.S.A.
REMARKS.-For description of skeleton see Auffenberg (1962d).

?Stylemys fbotti Stefano
?Stylemys bottii Stefano 1902a, p. 72.
Stylemys botti Williams 1950a, 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.

?Stylemys fcalaverensis Sinclair
?Stylemys calaverensis 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 \capax Hay
Stylemys nebrascensis Cope 1885, p. 770 ("No. 2") (part).
Stylemys capax 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 fconspecta 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


1974







BULLETIN FLORIDA STATE MUSEUM


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 fkarakolensis Riabinin
Stylemys karakolensis 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 fligonia (Cope)
Testudo ligonius Cope 1873a, p. 6.
Testudo ligonia Cope 1873b, p. 19.
?Stylemys ligonia Auffenberg 1962d, p. 8 [sic].
Geochelone 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 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.
GEOLOGIC RANGE.-Oligocene.
GEOGRAPHIC RANGE.-Northeastern Colorado, U.S.A.
REMARKS.-Probably conspecific with S. amphithorax (see Auffen-
berg 1962d).

Stylemys tnebrascensis Leidy
Emys hemispherica Leidy 1851a, p. 173.
Emys hemispherica Leidy 1851a, p. 173.
Testudo lata Leidy 1851a, p. 173.
Emys oweni Leidy 1851a, p. 327.
Emys culbertsonii Leidy 1852a, p. 34.
Testudo hemispherica Leidy 1852a, p. 59.
Testudo nebrascensis Leidy 1852a, p. 59.
Testudo oweni Leidy 1852a, p. 59.
Testudo culbertsonii Leidy 1852a, p. 59.
Emys nebrascensis Leidy 1852b, p. 65.
Emys lata Leidy 1852b, p. 65.
Stylemys culbertsonii Cope 1870, p. 124.


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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 fuintensis (Gilmore)
Testudo uintensis Gilmore 1915, p. 150.
?Stylemys uintensis Auffenberg 1962d, p. 10.
TYPE.-Carnegie Museum; a complete shell.
TYPE LOCALITY AND HoRIZON.-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. buresii).
Peltastes Gray 1869, p. 171 (Type: T. graeca L.).
Chersinella Gray 1870b, p. 725 (Type: T. graeca L.).
Testudinella Gray 1870b, p. 658 (Type: T. horsfieldii Gray).
Peltonia Gray 1872a, p. 4 (Type: T. graeca L.).
Medaestia Wussow 1916, p. 170 (Type: T. graeca L.).
Stylemys Auffenberg 1964d, p. 121 (part).
Agrionemys Khozatsky and Mlynarski 1966, p. 123 (part) (Type: T. horsfieldi Gray).
Agrione Khozatsky and Mlynarski 1966, p. 123 (typographical error).
Protestudo Chkikvadze 1970c, p. 7 (part) (Type: T. bessarabica).
GENOTYPE.-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.


1974







BULLETIN FLORIDA STATE MUSEUM


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 1870b, p. 658.
Agrionemys Khozatsky and Mlynarski 1966, p. 123 (as genus).
Agrione Khozatsky and Mlynarski 1966, p. 123 (typographical error).
TYPE SPECIEs.-Testudo horsfieldi 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) horsfieldi Gray
Testudo geometrica Pallas 1811, p. 19 (part).
Testudo horsfieldii Gray 1844, p. 74 (Afghanistan).
Chersus iberus Brandt 1852, p. 331 (part).
Homopus burnesii Blyth 1854, p. 642 (Afghanistan).
Testudinella horsfieldii Gray 1870b, 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


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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 horsfieldi will eventually be found to in-
habit much of the foothill area of the Himalaya Mountains. The recent
erection of Agrionemys for reception of horsfieldi (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 1870b, 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 graeca group according to Loveridge and Williams (1957).
REFERENCEs.-Skeleton: Siebenrock 1906, Williams 1950b, 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, fig. 4 (part).
Stylemys Auffenberg 1964d, p. 121 (part).
TYPE SPECIEs.-Testudo graeca Linnaeus (=Testudo [Testudo]
graeca Linnaeus).
DEFINITION.-A Eurasian 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;


1974








BULLETIN FLORIDA STATE MUSEUM


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 falba (Chkikvadze)
Protestudo alba Chkikvadze 1971, p. 245, fig. 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 (Testudo) fantiqua 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 neoviciensis Koch 1904, p. 60 (part) (typographical error).
Testudo antiqua var. noviciensis Glaessner 1933, p. 364.
Testudo antiqua noviciensis 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 praeceps Glaessner 1935, p. 125.
Testudo craverii Portis 1879, p. 121 [sic] (Sarmatian Miocene, Italy).
Testudo craver Pantanelli 1893, p. 135.
Testudo olawari 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 minute Fraas 1870, p. 51 (in error).


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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).
Stylemys canetotiana Auffenberg 1964d, p. 121.
Testudo csakvarensis Szalai 1934, p. 119 (Sarmatien Miocene, France).
Testudo csakvariensis Mlynarski 1966a (typographical error).
TYPE.-Fiirsten von Fiirstenberg? 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 antiqua 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) fbosporica 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 RANGE.-Crimea.
REMARKs.-Close to T. hermanni or T. graeca according to Riabinin
(1945); in the antiqua-graeca phyletic line according to Khozatsky
(1947).







BULLETIN FLORIDA STATE MUSEUM


Testudo (Testudo) catalaunica Bataller
Testudo catalaunica Bataller 1926, p. 149.
TYPE.-Seminario Councilar de Barcelona (Spain); a shell.
TYPE LOCALITY AND HORIZON.-St. Quirce, near Barcelona, 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 morphotype, not a subspecies, and therefore has no
validity as a trinomial. It is from the same locality as the type.

Testudo (Testudo) fcelonica Bergounioux
Testudo celonica Bergounioux 1958, p. 171; figs. 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 RANGE.-Late Miocene.
GEOGRAPHIC RANGE.-Northeastern Spain.
REMARKS.---Probably a synonym of T. antiqua.

Testudo (Testudo) tcernovi Khozatsky
Testudo cernovi 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 (Mlynarski
1968).
Testudo (Testudo) fcorroyi Bergounioux
Testudo corroyi 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.


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


GEOLOGIC RANGE.-Middle Eocene.
GEOGRAPHIC RANGE.-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 fdarewskii (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) fdenizoti Bergounioux
Testudo denizoti Bergounioux 1935, p. 87, pl. 5, fig. 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 1831b, 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) gglobosa Portis
Testudo globosa Portis 1890, p. 3.








BULLETIN FLORIDA STATE MUSEUM


TYPE.-Firenze Museum; a shell.
TYPE LOCALITY AND HoRIzoN.-Valley of the Arno 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 [Testudo] graeca terrestris Forskal,
Lebanon Mts., Israel, by Wermuth 1958) (not Testudo terrestris of Fermin
1765).
Testudo ibera Pallas 1814, p. 18 (=Testudo [Testudo] graeca ibera 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 1831a, 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, p. 309 (not of Schoepff).
Testudo ibera Gervais 1836, p. 309 (not of Pallas).
Testudo white Bennett 1840, p. 361 (In White).
Testudo graeca var. mauritanica Schlegel 1841, p. 106.
Testudo pusilla Strauch 1862, p. 67 (Transcaucasia) (part).
Peltastes marginatus var. white Gray 1870c, p. 11.
Chersinella graeca Gray 1873b, p. 725, pl. 60, fig. 4.
Testudo zarudyni Nikolsky 1869, p. 369 (=Testudo [Testudo] graeca zarudnyi
Nikolsky, Birdschau Prov., Eastern Iran, by Mertens 1946).
TYPE.-Location unknown to author.
TYPE LOCALITY.-Santa Cruz, Oran, Algeria, Africa (by Mertens and
Miiller 1928).
GEOLOGIC RANGE.-Pleistocene of Morocco (Lecointre 1926), France
(Henri-Martin 1946) and Recent.
GEOGRAPHIC RANGE.-Northwestern Africa from Morocco and Malta
to Cyrenaica region of Libya.
REMARKS.-See 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,


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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 1950b, 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) fcanstadiensis 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 enriquesi 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) ihipparionum Wiman
Testudo hipparionum Wiman 1930, p. 41, pl. 6, fig. 2-2b.
Geochelone 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







BULLETIN FLORIDA STATE MUSEUM


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) fhonanensis 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 filliberalis (Chkikvadze)
Protestudo illiberalis Chkikvadze 1971, p. 246, fig. 3.
TYPE.-Georgian Academy of Science (Tbilisi); nearly complete
plastron imbedded in nodule.
TYPE LOCALITY 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 horsfieldi.

Testudo (Testudo) tkegenika Khozatsky
Testudo kegenika Khozatsky 1955, p. 51.


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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 RANGE.-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) kucurganica 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 LOCALITY AND HORIzoN.Novopetrovka, Alekseeva and other
localities 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) flamanoni 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 vidali.

Testudo (Testudo) lunellensis Almera and Bofill
Testudo lunellensis Almera and Bofill 1903, p. 106 [sic].
Testudo lunellensis Deperet 1906, p. 12.








BULLETIN FLORIDA STATE MUSEUM


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 praegraeca 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 LOCALITY 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 tabulata var. campanulata Walbaum 1782, p. 124 (nomen illegitum).
Testudo marginata Schoepff 1792, p. 58, pls. 11-12, fig. 1 (unknown).
Testudo graja 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).
Peltastes marginatus Gray 1869, p. 173.
Testudo nemoralis Schreiber 1875, p. 557 (Greece).
TYPE.-Location unknown to author.
TYPE LOCALITY-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.


Vol. 18, No. 3







AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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
1950b, Auffenberg 1966b; Hyobranchial Apparatus: Siebenrock 1898;
Skull: Siebenrock 1897; Serology: Obst and Ambrosius 1971.

Testudo (Testudo) f marmorum Gaudry
Testudo marmorum 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) foriens 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 LOCALITY 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.







BULLETIN FLORIDA STATE MUSEUM


Testudo (Testudo) froguesi 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.-Close 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) tseminota 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.-C1OSe to hermanni-graeca.


Testudo (Testudo) \serresi Pictet
Testudo serresii Pictet 1845, p. 20.
Testudo serresi Bergounioux 1938a, p. 279.
TYPE.-Location unknown to author; a shell.
TYPE LOCALITY 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).


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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 Paleoanthropology
(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) Isphaerica Wiman
Testudo sphaerica Wiman 1930, p. 33, figs. 2-26.
Terrapene sinica Young 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)
horsfieldi. 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. yushensis Yeh is placed here in view of its
similarity in morphology and geologic age.

Testudo (Testudo) isuttoensis 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 LOCALITY AND HORIZON.-Sutto, Komarom County, Hungary;
Middle Pleistocene.







BULLETIN FLORIDA STATE MUSEUM


GEOLOGIC RANGE.-Middle Pleistocene.
GEOGRAPHIC RANGE.-Hungary.
REMARKS.-A poorly defined form close to T. marginata (Szalai
1934) and in the "graeca-group" (Szalai 1936).

Testudo (Testudo) fszalai 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) ftunhuanensis 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).


Vol. 18, No. 3








AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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, Tarn Department,
France; Ludian faunal age, 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.-For a discussion of relationships see Loveridge and Wil-
liams (1957), who consider it close to T. scutella.

?Testudo jptychogastroides Reinach
Testudo ptychogastroides Reinach 1900, p. 19.
TYPE.-?Frankfurt Museum, Germany; shell fragment.


1974







BULLETIN FLORIDA STATE MUSEUM


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 freidlii Hoernes
Testudo reidlii Hoernes 1892, p. 243.
"Testudo"(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 fscutella (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 jstehlini 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 Brim (1951) for best description; if his statement is
correct that stehlini is closely related to pyrenaica, then it probably be-
longs in the genus Geochelone.


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


?Testudo tungia Yeh
Testudo tungia Yeh 1963b, p. 224, fig. 1, pl. 1.
TYPE.-Institute of Vertebrate Paleontology and Paleoanthropology
(Peking); a carapace.
TYPE LOCALITY AND HORIZON.-Gigantopithecus Cave, Liucheng,
Kwangsi, China; Lower Pleistocene.
GEOLOGIC RANGE.-Pleistocene.
GEOGRAPHIC RANGE.-Kwangsi, China.
REMARKS.-PrObably not even a testudinid; perhaps referable to the
genus Cuora.

Genera Inquirendae

Genus fCheirogaster Bergounioux
Cheirogaster Bergounioux 1935, p. 78.
GENoTYPE.-Cheirogaster maurini Bergounioux (by monotypy).
DEFINITION.-If correctly described, this is a monotypic genus char-
acterized by lack of entoplastron and contact between gular and pectoral
scutes.
TYPE LOCALITY AND HomnzoN.-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-
examination 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 imaurini 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 (Preliminary notice).
Cheirogaster arrahonensis Bergounioux 1958, p. 175, figs. 15, 16, pis. 32, 33.







BULLETIN FLORIDA STATE MUSEUM


TYPE.-Museo de Sabadell (Spain); a partial shell.
TYPE LOCALITY AND HORIZON.-Sabadell, Penedes District, Spain;
Vallesian faunal age, Late Miocene.
GEOLOGIC RANGE.-Late Miocene.
GEOGRAPHIC RANGE.-Northeastern Spain.
REMARKS.-Very poorly defined.

Genus fFloridemys Williams
Bystra Hay 1916a, p. 53 (preoccupied by Bystra Cameron 1902).
Floridemys Williams 1950a, p. 27 (substitute name).
GENOTYPE.-Bystra nanus Hay ( = Floridemys nana [Hay] [by mon-
otypy]).
DEFINITION.-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 HomzON.-Holder Phosphate Mine, near Inver-
ness, Citrus County, Florida, U.S.A.; Hawthorne Formation, Miocene
(Early?).
GEOLOGIC RANGE.-Miocene (Early?).
GEOGRAPHIC RANGE.-Central Florida, U.S.A.
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 tKansuchelys Yeh
Kansuchelys Yeh 1963a, p. 28.
GENOTYPE.-Kansuchelys chiayukuanensis Yeh.
DEFINITION.-Small to medium tortoises with hexagonal neurals
throughout the series, and a doubly notched epiplastral projection.


Vol. 18, No. 3








AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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 tchiayukuanensis Yeh
Kansuchelys chiayukuanensis 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 fovalis Yeh
Kansuchelys ovalis 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 tSinohadrianus Ping
Sinohadrianus Ping 1929, p. 232.
GENOTYPE.-Sinohadrianus sichuanensis Ping.
DEFINITION.-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.








BULLETIN FLORIDA STATE MUSEUM


Sinohadrianus teozensis Shikama
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 fsichuanensis 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
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Stylemys nebrascensis Leidy. Copeia, 1961 (4): 496-498.


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AUFFENBERG: FOSSIL TORTOISE CHECKLIST


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Zoological Society, 7: 1-17.


1974








BULLETIN FLORIDA STATE MUSEUM


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1974




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