0876669186

Free ebooks ==> www.Ebook777.
com
www.Ebook777.com
Unless otherwise credited, all photographs are by the author.
ISBN 0-87666-918-6
© 1979 T.F.H. Publications, Inc. Ltd.
Distributed in the U.S. by T.F.H. Publications, Inc., 211 West

Sylvania Avenue, P.O. Box 427, Neptune, N.J. 07753; in
England by T.F.H. (Gt. Britain) Ltd., 13 Nutley Lane, Reigate,
Surrey; in Canada to the book store and library trade by Beaver-
books, 953 Dillingham Road, Pickering, Ontario L1W 1Z7; in
Canada to the pet trade by Rolf C. Hagen Ltd., 3225 Sartelon
Street, Montreal 382, Quebec; in Southeast Asia by Y.W. Ong, 9
Lorong 36 Geylang, Singapore 14; in Australia and the South
Pacific by Pet Imports Pty. Ltd., P.O. Box 149, Brookvale 2100,
N.S.W., Australia; in South Africa by Valiant Publishers (Pty.)
Ltd., P.O. Box 78236, Sandton City, 2146, South Africa; Publish
ed by T.F.H. Publications, Inc., Ltd., The British Crown Colony
of Hong Kong.
Free ebooks ==> www.Ebook777.com
Contents
I ’reface........................................................................................ 5
I TURTLE IDENTIFICATION.............................................................. 7
2. TURTLE STRUCTURE AND FUNCTION......................................... 18
.1 TURTLE EVOLUTION AND FOSSIL HISTORY.................73
■I EMYDID TURTLES......................................................................108
Pseudemys - 110; Chrysemys - 135; Graptemys -
137; Malaclemys - 150; Deirochelys - 154; Emy-
doidea - 157; Clemmys - 158; Terrapene - 163;
Rhinoclemys - 172; Emys - 183; Mauremys - 185;
Sacalia - 190; Annamemys - 191; Hardella - 193;
Iiatagur - 194; Callagur - 196; Orlitia - 197;
Kachuga - 198; Geoemyda - 201; Heosemys - 201;
Melanochelys - 204; Cyclemys - 206; Cuora - 211;
Hieremys - 213; Notochelys - 215; Ocadia - 216;
Morenia - 217; Geoclemys - 218; Chinemys - 219;
Malayemys - 223; Siebenrockiella - 224; Back
ground Reading - 226.
5 LAND TORTOISES....................................................................... 247
Geochelone - 258; G. (Asterochelys) - 275; G.
(Aldabrachelys) - 286; G. (Cylindraspis) - 302;
G. (Manouria) - 311; G. (Indotestudo) - 314; G.
(Chelonoidis) - 322; Testudo - 394; Gopherus -
418; Psammobates - 438; Homopus - 449; Kin-
ixys - 454; Chersine - 461; Malacochersus - 463;
Pyxis - 466; Acinixys - 468; Background Read
ing - 469.
(I. MUD, MUSK, AND SNAPPING TURTLES..........................................481
Chelydra - 481; Macroclemys - 493; Kinosternon -
505; Sternotherus - 557; Staurotypus - 572;
Claudius - 580; Background Reading - 589.
7. SOFT-SHELLED TURTLES.............................................................. 600
IAssemys - 608; Cyclanorbis - 613; Cycloderma -
617; Trionyx - 621; Chitra - 650; Pelochelys - 652;
Background Reading - 653.
www.Ebook777.com
8. MONOTYPIC TURTLE FAMILIES..................................................656
Carettochelys - 656; Dermatemys - 661; Platy-
sternon - 666; Background Reading - 670.
9. SEA TURTLES............................................................................. 672
Chelonia - 676; Caretta - 693; Lepidochelys - 701;
Eretmochelys - 713; Dermochelys - 722; Back
ground Reading - 732.
10. SIDE-NECKED TURTLES............................................................ 748
Podocnemis - 750; Peltocephalus - 758; Erym-
nochelys - 760; Pelusios - 761; Pelomedusa - 770;
Chelus - 772; Hydromedusa - 775; Platemys - 778;
Phrynops (Mesoclemmys) - 782; P. (Phrynops) -
784; P. (Batrachemys) - 789; Chelodina - 795;
Emydura - 803; Elseya - 807; Pseudemydura -
810; Background Reading - 813.
11. TURTLE CONSERVATION AND EXPLOITATION. . .818
12. TURTLES IN CAPTIVITY..............................................................833
APPENDIX I. CHECKLIST................................................................... 850
APPENDIX II. TURTLE GENERA..........................................................860
GLOSSARY......................................................................................... 876
ABOUT THE AUTHOR
INDEX
PREFACE
The original intention in writing this book was to provide an
updated and revised version of my 1967 book Living Turtles of the
World. However, at an early stage in the work it became clear that
not only my personal knowledge of turtles, but also the sum total of
human knowledge of these animals, had advanced to such an ex
tent that it would be more appropriate to write a completely new
book. The advances in my personal knowledge during the last
decade have been made possible by the opportunities presented by
my graduate work at the University of Florida, where I had the
privilege of working with Dr. Archie Carr, and my subsequent em
ployment as Co-ordinator of the Marine Turtle Group of the Inter
nation Union for the Conservation of Nature and the World Wildlife
Fund. To these institutions I am profoundly thankful. The present
work, then, will be reminiscent of Living Turtles in some of its pho
tographs and chapter headings, but beyond that the similarity
more or less ends, and we feel justified in using the new title.
It is, perhaps, a presumptuous task for one whose field work
has been nearly (though not completely) confined to the New
World to attempt an even and thorough coverage of all of the tur
tles of the world. Indeed, to a grea,t extent the available informa
tion on living turtle species is so uneven that the task may be con
sidered impossible. The presumption is perhaps heightened by the
ambition of this book to be all things to all men—or at least to all
turtle men; I have attempted to write a book which will be of value
to pet keepers, scientific turtle experts, and everyone in between,
being particularly mindful of the intelligent general naturalist and
the academic zoologist who would have use for an overview of the
Testudines within the covers of a single volume. I have felt justi
fied in undertaking this presumptuous task partly because there is
need for such a book, partly because regional studies by many
gifted cheloniophiles in recent years have opened up enormous
amounts of new information, and partly because my own burning
interest in turtles has made the writing of this book, despite the
long hours of research and typing, a labor of love.
The theme of Encyclopedia of Turtles is the diversity of turtles
and their many wonderful adaptations for different ways of life.
The range of turtle habitats, from the open ocean to African rock
crevices to prairies, bogs, ponds, and open lakes, is surely at least
5
www.Ebook777.com
as great as that of any other animal group of similar size, and tur
tles have responded to this diversity of habitat by evolving a fasci
nating variety of shapes and sizes. On the other hand, there is little
mention in this book of the laboratory and experimental aspects of
the study of turtles. This is not because I deem such studies less
important, but because they will be handled much more thorough
ly than I ever could in a forthcoming volume to be edited by Henry
Morlock and Marion Harless, for which I have been honored by
being asked to write the opening chapter.
A weakness of Living Turtles of the World was its scant cover
age of the literature. I have attempted to rectify this in the present
work, not by means of frequent textual citations which I felt would
be inappropriate in a work of this nature and would make for inter
rupted reading, but by providing a “background reading” section
at the end of each chapter. It has, of course, been an invidious task
to select the nearly 1000 papers and books cited (with some dupli
cation) from the enormous volume of turtle literature now avail
able. Men have been fascinated by turtles for centuries, and, for
better or worse, most of those who have been fascinated by them
have felt driven to record their thoughts and observations for pos
terity. I defend only the commission aspect of the works cited—I
believe each of them will repay the reader’s time in finding and
reading it. I am aware that I must have omitted many important
papers completely; to the authors of same, I apologize, and intend
no slight. However, the literature that is cited is sufficient to pro
vide a very thorough coverage of present knowledge of turtles,
while if we include “second generation” references—those listed
in the publications cited in this book—coverage of the literature
would be incomplete only in the omission of those works which
were published too recently to be included.
6
Chapter 1
TURTLE IDENTIFICATION
Since this is the only book in English which describes all living
turtle species, it is inevitable that one of its main uses will be for
identification, particularly of turtles whose origin is unknown. The
normal way to do this, and usually the quickest, is to scan the pho
tographs and, having found one which bears a reasonable resemb
lance to the specimen in hand, to refer to the appropriate section of
the text. Nevertheless, this somewhat unsystematic method does
have shortcomings; unrelated turtles can, by parallelism, have a
similar superficial appearance, and atypical specimens of
familiar species can sometimes look very different from the norm.
For this reason, taxonomists spend much of their time drawing up
‘keys’ for systematic identification of specimens. The most usual
type is the dichotomous key, in which the user is led through pairs
of alternative characterizations; thus, the specimen in hand may
either correspond to alternative 1, or to alternative 1’. After each
characterization, the user is referred either to a subsequent pair of
alternatives or is presented with the final identification of the spe
cimen.
In theory, one cannot go wrong when following a properly de
signed key. In practice, however, there are a number of short
comings, particularly in a key for identification of genera rather
than of species within a genus. This is because a genus can be,
superficially, much less distinctive than a species. For example,
the spotted turtle, Clemmys guttata, is one of the most distinctive
and readily identified of all turtles. On the other hand, to identify
the genus Clemmys and differentiate it from other emydid genera,
one has to examine the skull, the structure of the jaw surfaces, and
the shape of the neural bones—characters which require skeletal
material for sure checking. Another difficulty is that taxonomists,
in their striving for absolute accuracy in keys, may pass over easi
ly checked, external characters which separate the forms con
cerned in almost all cases, and refer instead to minor, internal
characters which will always hold up. Moreover, these minor in
ternal characters may be impossible to express in common lang
uage, and the student who has reached the point of understanding
all the technical terms employed will probably also be familiar
7
www.Ebook777.com
with the various genera and species in the group, and hence have
no need for a key.
Despite these shortcomings, dichotomous identification keys
have their uses, and so a key to turtle families, and to the genera of
each of the polytypic families, is given below. By scanning the pic
tures and text in the book, and by using the keys as far as possible,
the student should eventually be able to identify any turtle that he
may encounter. However, to reach that happy point where he can
identify a turtle at sight, even if only an empty shell or otherwise
incomplete specimen is available, takes years of experience in
handling and examining living and preserved turtles of all kinds,
and of reading widely and deeply in the field.
It might be mentioned parenthetically that keys are of two
kinds, artificial and natural, though for straightforward identifica
tion purposes the methods of using them, and the results obtained,
are similar. Artificial keys may be easier to use, however, since
they can utilize the most conspicuous, rather than the most funda
mental characters. For example, in an artificial key to the genera
within a certain turtle family, a monotypic genus could be identi
fied by some striking color pattern characteristic of the only spe
cies included in that genus, rather than by some internal but more
fundamental character that characterizes the genus itself. On the
other hand, natural keys have two advantages: they enable the
user to make a correct generic identification even of a specimen of
a new, undescribed species of a known genus; and they give the
user some indication of how the genera within a family are actual
ly defined. A result of this is that in a natural key only one combi
nation of alternatives will lead to the identification of a particular
genus, while in an artificial key several different combinations of
characters, or several different paths through the key, may lead to
the same genus. Thus a natural key will have one couplet fewer
than the total number of taxa identified, while an artificial key
may have many more couplets than the number of taxa concerned.
The keys given below are natural keys. The reader who wishes to
compare the usage of a natural key with that of an artificial key
may be interested In using the natural key to the genera of the
Emydidae given below with the artificial key to these genera given
by Wermuth and Mertens (1961: Schildkroten, Krokodile, Bruche-
nechsen; Gustav Fischer, Jena; pp. 40-45). It is noteworthy that
the key of Wermuth and Mertens, which recognizes 25 genera, has
49 couplets, while the one used here, which recognizes 31 genera
and subgenera (the greater number is due to the fragmentation of
the old genera Clemmys and Geoemyda), has only 30 couplets.
Nevertheless, no value judgement is implicit here, and it is quite
possible that most readers will find the key of Wermuth and Mer
tens easier to use.
8
KEY TO TURTLE FAMILIES

1 Carapace and plastron covered with clearly demarcated homy
scutes............................................................................... 2
1’ Carapace and plastron covered with a continuous layer of un
divided skin.................................................................... 10
2 Front limbs in form of elongate flippers, without separate
digits; each forelimb with one or two claws; bony skull roof
complete and intergular scute present..............Cheloniidae
2’ Front limbs with distinct digits or club shaped, each bearing
four or five claws; bony skull roof usually emarginate;
intergular present or absent..............................................3
3 Neck retracted in horizontal plane; pelvis fused to shell; inter
gular scute present........................................................... 4
3’ Neck retracted in vertical plane; pelvis not fused to shell;
intergular scute absent..................................................... 5
4 Skull roof usually emarginate from behind, not or but slightly
emarginate from below; mesoplastra present........................
................................................................... Pelomedusidae
4’ Skull roof emarginate from below, not or but slightly from be
hind; mesoplastra absent...................................... Chelidae
5 Entoplastron present or absent and front part of plastron
moveable about a hinge between epiplastral and hyoplastral
bones (between pectoral and abdominal scutes), or ento
plastron present and plastron rigid but composed of only
eight scutes and attached to carapace by a flexible and very
narrow bridge................................................ Kinostemidae
5’ Entoplastron present; plastron, if hinged, with hinge situated
between hyo- and hypoplastral bones (or between epiplas
tral and hyoplastral bones and humeral and pectoral scutes
in Pyxis, between abdominal and femoral scutes in some fe
male Testudo)\ more than eight plastral scutes present;
bridge not both very narrow and flexible .......................... 6
6 One or more inframarginal scutes present between axillary
and inguinal scutes on each side of plastron, separating pec
toral and abdominal scutes from marginal scutes............. 7
6’ No inframarginal scutes present, pectoral and/or abdominal
scutes in contact with marginal scutes ............................ 9
7 Tail short, head small and jaws denticulate and unhooked,
plastron large, skull roof emarginate from behind................
.................................................................. Dermatemyidae
7’ Tail long and armored, head very large and with non-denticu-
late, hooked jaws; either plastron reduced and cruciform
but with rigid bridges and skull roof emarginate, or plastron
moderately large but with flexible bridges and skull roof
complete........................................................................... 8
www.Ebook777.com 9
8 Plastron reduced and cruciform, rounded anteriorly, bridges
rigid, abdominal scutes widely separated on midline, skull
roof emarginate................................................. Chelydridae
8’ Plastron moderately large, truncated anteriorly, bridges flex
ible, abdominal scutes with median line of contact, skull roof
complete....................................................... Platystemidae
9 Habitat terrestrial, hind feet elephantine, osteoderms usually
present in forelimbs, never more than two phalanges in each
digit of hind foot...............................................Testudinidae
9’ Habitat typically aquatic, hind feet not elephantine, no osteo
derms in forelimbs, always more than two phalanges in at
least one digit of hind foot.................................... Emydidae
10 Carapace raised into seven prominent longitudinal ridges;
upper jaw strongly cusped; limbs clawless . .Dermochelyidae
10’ Carapace not ridged; upper jaw not strongly cusped; claws
present .............................................................................11
11 Hind part of carapace flexible; plastral bones reduced.............
....................................................................... Trionychidae
11’ Entire carapace rigid, plastron completely ossified...................
.................................................................Carettochelyidae
KEYS TO TURTLE GENERA

No keys are required for the Dermochelyidae, Der-
matemyidae, Platystemidae, and Carettochelyidae, since each of
these families includes only one genus.
KEY TO THE CHELONIIDAE

1 Four costal scutes on each side of carapace........................... 2
1’ Five or more costal scutes on each side of carapace................3
2 Scutes overlapping (except in hatchlings and old adults), head
very narrow, jaw surfaces not serrated, two pairs of pre
frontal scales.................................................. Eretmochelys
2’ Scutes juxtaposed, head not particularly narrow, rounded
anteriorly; jaw surfaces serrated, one pair of prefrontal
scales.................................................................... Chelonia
3 Overall coloration red-brown, shell rather elongate, no pores in
inframarginal scutes................................................Caretta
3’ Overall color grey- or olive-green, shell very broad, inframargi
nal pores present............................................ Lepidochelys
KEY TO THE PELOMEDUSIDAE

1 Front lobe of plastron attached by flexible hinge; mesoplastra
with median line of contact..................................... Pelusios
1’ Plastron rigid, mesoplastra well separated................................2
10
2 Hind feet with five claws; shell bones thin, with large fontanelle
in middle of plastron; skull roof fully emarginate.................
........................................................................ Pelomedusa
2’ Hind limbs with four claws; shell bones thick, without mid-
plastral fontanelle except in very young; skull roof nearly
complete..............................................................................3
3 Longitudinal groove present between eyes; orbits oriented
dorsally on skull; upper jaw notched or rounded off but not
hooked...............................................................Podocnemis
3’ No groove between eyes; orbits laterally oriented on skull;
upper jaw slightly or strongly hooked.................................4
4 Intergular scute short, not separating gulars; skull broad, not
elongate anteriorly; no premaxillary fossa.............................
......................................................................Erymnochelys
4’ Intergular scute long, separating gulars; skull triangular,
broad and deep posteriorly, somewhat elongated anteriorly;
a deep premaxillary fossa present................... Peltocephalus
KEY TO THE CHELIDAE

1 Head very flat, decorated with numerous fleshy tassels and ex
crescences ; tip of snout produced into snorkel-like breathing
tube..........................................................................Chelus
1’ Head not so flat, without excrescences except for simple
barbels on chin; tip of snout not forming breathing tube . . . . 2
2 Neck strikingly elongate, nearly as long as shell .................... 3
2’ Neck much shorter than shell ................................................4
3 Nuchal scute large but excluded from front margin of carapace
by mesially contiguous first marginals; temporal arch pres
ent; neural bones present; intergular scute completely
separating gulars ..........................................Hydromedusa
3’ Nuchal scute not excluded from front margin of carapace; no
temporal arch; neural bones usually absent; intergular scute
not completely separating gulars.......................... Chelodina
4 Intergular scute very large, completely separating gulars and
humerals and partially separating pectorals; skull roof only
slightly emarginate ......................................Pseudemydura
4’ Intergular scute smaller, not completely separating humerals;
skull roof extensively emarginate ...................................... 5
5 First vertebral not strikingly wide (narrower than the second
vertebral); symphysis of lower jaw broader than diameter
of orbit............................................................................... 6
5’ First vertebral strikingly wide, wider than the second verte
bral; symphysis of lower jaw narrower than the diameter of
the orbit............................................................................ 7
6 Crown of head covered with a single, distinct, homy shield;
erect, pointed tubercles on back of neck................... Elseya
11
www.Ebook777.com
6’ Crown of head not covered by a single homy cap; blunt papillae
on top of neck....................................................... Emydura
7 Carapace with a distinct median groove..................... Platemys
7’ Carapace without a distinct median groove.......................................... 8
8 Head very large (width of head about (4 length of carapace);
adult size large—ten to eighteen inches.............................. 9
8’ Head not very large (width about 1/6 length of carapace); adult
size small—maximum length about seven inches.................
.................................................. Phrynops (Mesoclemmys)
9 Parietal-squamosal arch very slender; parietal bones not ex
panded dorsally....................... . .Phrynops (Batrachemys)
9’ Parietal-squamosal arch stronger; parietal bones expanded
dorsally, producing a flat bony area between the temporal
muscles..............................................Phrynops (Phrynops)
KEY TO THE KINOSTERNIDAE

1 Entoplastron absent, usually eleven (sometimes ten) plastral
scutes present.................................................................... ..
1’ Entoplastron present, usually eight (sometimes 9) plastral
scutes present.................................................................... ..
2 Plastron large or slightly reduced, interpectoral seam usually
very short; plastron rounded anteriorly; two fully functional
plastral hinges in adults....................................Kinostemon
2’ Plastron reduced, often with areas of skin between plastral
scutes; interpectoral seam not very short; plastron some
what truncated anteriorly, and with a well-developed anteri
or hinge but rather feeble posterior hinge in adults..............
.......................................................................Stemotherus
3 Carapace with three very strong longitudinal keels (especially
in adults); anterior lobe of plastron hinged; bridges rigid;
nine plastral bones...........................................Staurotypus
3’ Carapace smooth or with only feeble indications of longitudinal
keels; no plastral hinge; bridges flexible; seven plastral
b°nes...................................................................... Claudius
KEY TO THE CHELYDRIDAE

1 Supramarginal scutes present; eyes on side of head; worm-like
fishing lure on base of mouth.........................Macroclemys
1’ Supramarginal scutes absent; eyes inclined toward top of head;
no lure on floor of mouth.......................................Chelydra
KEY TO THE TESTUDINIDAE

1 Front lobe of plastron moveable about hinge between humeral
and pectoral scutes..................................................... Pyxis
12
1' Front lobe of plastron not moveable ...................................... 2

2 Carapace exceedingly flat, fenestrated and flexible throughout
life................................................................ Malacochersus
2’ Carapace more or less domed, not fenestrated or flexible in
adults................................................................................ 3
3 Hind part of carapace moveable about a hinge between the
second and third costals and the seventh and eighth margi
nals; outer margin of 4th costal markedly wider than outer
margin of 3rd; hind margin of plastron truncated ... Kinixys
3’ Entire carapace rigid, without hinge; outer margin of 3rd costal
wider than outer margin of 4th; plastron not truncated, fre
quently notched ................................................................ 4
4 Median ridge on premaxilla; forelimbs flattened, adapted for
digging................................................................. Gopherus
4’ No median ridge on premaxilla; forelimbs not adapted for
digging ..............................................................................5
5 Gular scute single, projecting, truncated anteriorly ... Chersina
5’ Gulars normally paired, or if single, not truncated anteriorly.. 6
6 At least some of the neural bones alternately quadrilateral and
octagonal; upper ends of pleural bones alternately wide and
narrow............................................................................... 7
6’ Neural bones hexagonal, upper ends of pleural bones not alter
nately wide and narrow..................................................... 8
7 Supranasal scales present; nuchal scute present; not more than
one tubercle on each thigh; suprapygal single or divided
transversely; posterior lobe of plastron in adults at least
somewhat moveable in one or both sexes; prootic usually
concealed dorsally by the parietals..........................Testudo
7’ Supranasal scales absent; nuchal scute present or absent; up to
three tubercles on each thigh; two suprapygals, the anterior
one embracing the smaller posterior one; posterior lobe of
plastron always rigid; prootic usually well exposed dorsally
............................................................. Geochelone
8 Maxilla with triturating ridge; dorsal surface of tail covered
with flattened scale................................................ Acinixys
8’ Maxilla unridged; dorsal surface of tail not covered with flat
tened scale .......................................................................®
9 Carapace somewhat depressed; vertebral scutes never conical;
each gular usually broader than long; anterior palatine fora
men large, conspicuous; maxillary bone not entering roof of
palate .................................................................. Homopus
9’ Carapace rather convex; vertebral scutes somewhat conical;
each gular usually as long as or longer than broad; anterior
palatine foramen small, concealed; maxillary bone entering
roof of palate..................................................Psammobates
13
www.Ebook777.com
KEY TO THE EMYDIDAE
1 Supracaudal scutes fall short of suture between pygal and
suprapygal; a double joint between fifth and sixth cervical
vertebrae; no strong lateral tuberosity on basioccipital;
angular bone forms floor of canal for Meckel’s cartilage;
New World turtles, except for Emys.................................... 2
1’ Supracaudal scutes extend forward on to suprapygal; a single
joint between fifth and sixth cervical vertebrae; a strong
lateral tuberosity on the basioccipital that forms the floor of
the recessus scalae tympani; angular bone excluded from
contact with Meckel’s cartilage by a longitudinal flange of
the articular; Old World, except for Rhinoclemys................ 10
2 Plastron hinged, at least in half-grown to adult specimens, be
tween the pectoral and abdominal scutes........................... 3
2’ Plastron rigid throughout life ................................................. 5
3 Shell openings completely closed when plastral lobes are
raised, except in very young specimens................Terrapene
3’ Shell openings not completely closeable at any age............... 4
4 Head and neck elongate; ribs bowed inward at point of attach
ment to vertebral column ..................................Emydoidea
4’ Head and neck not strikingly elongate; ribs not bowed inward
at point of attachment to vertebral column ................Emys
5 Alveolar surface of upper jaw with a ridge or row of tubercles
extending parallel to its margin.........................................6
5’ Alveolar surface of upper jaw smooth or undulating, not ridged
................................................................................................... 7
6 Carapace smooth, keel-less, hind margin not serrate...............
......................................................................... Chrysemys
6’ Carapace with longitudinal wrinkles, a more or less complete
keel, or both; hind margin somewhat serrate... .Pseudemys
7 Alveolar surface of upper jaw narrow, its inner edge parallel to
the cutting edge; carapace not tuberculate......................... 8
7’ Alveolar surface of upper jaw either very broad, forming most
of the roof of the anterior part of the mouth (a ‘false palate’),
or broadened posteriorly and much narrower toward the
symphysis; vertebral scutes tuberculate except in certain
old adults........................................................................... 9
8 Neck very long; ribs bowed inward at point of attachment to
vertebral column............................................... Deirochelys
8’ Neck not very long; ribs not bowed inward at point of attach
ment to vertebral column .....................................Clemmys
9 Scutes of the carapace usually with concentric ridges or stria-
tions; head and neck plain, spotted, or mottled, without
longitudinal light stripes................................... Malaclemys
9’ Scutes of the carapace smooth, not concentrically striated or
14
ridged; head and neck with longitudinal light stripes...........

..........................................................................Graptemys
10 A hinge present in the plastron between the humeral and pecto
ral scutes, at least in adults............................................11
10’ Plastron entirely rigid at all ages.........................................13
11 Hinges well-developed and plastral lobes large enough to close
shell openings completely; hind margin of carapace smooth
.................................................................................Cuora
11’ Plastral hinges only properly developed in adult specimens;
plastral lobes too small to allow shell openings to be com
pletely closed; hind margin of carapace serrate................. 12
12 Five vertebral scutes ................................................ Cyclemys
12’ Six (or seven) vertebral scutes.................................Notochelys
13 Triturating surfaces of upper jaw very broad, the upper sur
face with a pair of middle ridges that end anteriorly in a pair
of cusps.......................................................................... 14
13’ Triturating surfaces of upper jaw broad or narrow, but middle
ridges, if present, not ending anteriorly in a pair of cusps.. 16
14 Carotid canal between pterygoid and prootic; chamber for
paracapsular sac completely open posteriorly......... Hardella
14’ Carotid canal entirely within pterygoid; chamber for paracap
sular sac not completely open posteriorly........................15
15 Sculpture of triturating surfaces sharply defined; plastral but
tresses weak .........................................................Morenia
15’ Sculpture of triturating surfaces only faintly indicated on the
bone; plastral buttresses moderate................... Geoclemys
16 Triturating surfaces narrow and without bony ridging.......... 17
16’ Triturating surfaces broad, at least posteriorly, usually with
bony ridging .................................................................... 23
17 Entire horizontal temporal arch, including squamosal bone,
absent............................................................... Heosemys
17’ Squamosal bone present, though may be loosely attached ... 18
18 Inferior process of parietals convergent ventrally, encroach
ing on groove for trabecular cartilage, so that cranial cavity
is markedly narrowed ventrally......................... Geoemyda
18’ Inferior process of parietals not convergent ventrally........ 19
19 Hexagonal neurals usually short-sided in front; tail long in
young, moderate in adult................................................ 20
19’ Hexagonal neurals short-sided behind; tail short................22
20 Plastral buttresses very strong, nearly reaching neural bones
............................................................................ Annamemys
20’ Plastral buttresses not reaching near neural bones............ 21
21 Fissura ethmoidalis keyhole-shaped; forearm with numerous
transversely enlarged scales separated by granular scales;
plastral buttresses moderately strong............... Mauremys
21’ Fissura ethmoidalis broadly triangular, forearm covered with
15
www.Ebook777.com
a few very large scales without intervening granules; plas-
tral buttresses very weak.........................................Sacalia
22 Carapace with a median keel or unkeeled; ventral end of jugal
broad............................................................... Rhinoclemys
22’ Carapace tricarinate, ventral end of jugal narrow.....................
......................................................................... Melanochelys
23 Vomer reduced, its posterior end excluded from cranial cavity
by pterygoids; dentary symphysis extending well behind
lower triturating surfaces along midline.......................... 30
23’ Vomer large, its posterior end extending back to the level of
the cranial cavity; lower triturating surface equalling or
exceeding dentary symphysis in backward extent along the
midline............................................................................24
24 First hexagonal neural bone short-sided in front, those pos
terior to this short-sided behind......................................25
24’ Hexagonal neural bones short-sided in front ......................26
25 Bony temporal arch complete; entoplastron anterior to the
humero-pectoral suture.....................................Malayemys
25’ Temporal arch incomplete, entoplastron intersected by the
humero-pectoral suture........................................ Hieremys
26 Upper triturating surface without traces of ridging.. Chinemys
26’ Upper triturating surface with one or two ridges................... 27
27 Upper triturating surface with two denticulated ridges.............
............................................................................... Batagur
27’ Upper triturating surface with a single sharply-defined and
denticulated median ridge................................................. 28
28 Humero-pectoral sulcus traversing entoplastron; two phalan
ges in fifth digit; fourth vertebral not elongated; plastral
buttresses somewhat reduced.................................. Ocadia
28’ Humero-pectoral sulcus not traversing entoplastron; three
phalanges in fifth digit; fourth vertebral scute often elonga
ted ; plastral buttresses very strong................................ 29
29 Fourth central scute overlying parts of four or five neural
bones.....................................................................Kachuga
29’ Fourth central scute overlying parts of three neural bones.......
.............................................................................Callagur
30 Nasopalatine foramen large, exceeding the posterior palatine
foramen in size; upper triturating surfaces broad posterior
ly, with sharply defined middle ridge; neural bones elonga
ted; entoplastron anterior to humero-pectoral sulcus; plas
tral buttresses very strong......................................... Orlitia
30’ Nasopalatine foramen smaller than the posterior palatine for
amen ; triturating surfaces moderately narrow, without dis
tinct middle ridge; neural bones short and nearly square in
the middle of the series; humero-pectoral sulcus traversing
entoplastron; plastral buttresses moderate . Siebenrockiella
16
KEY TO THE TRIONYCHIDAE

1 Plastron with flexible flaps protecting the hind limbs; seven or
more plastral callosities.................................................... 2
1’ Plastron without flexible flaps protecting hind limbs; only four
plastral callosities............................................................. 4
2 Bony arcade between orbit and temporal emargination broader
than the diameter of the orbit........................... Cycloderma
2’ Bony arcade between orbit and temporal emargination nar
rower than diameter of the orbit ...................................... 3
3 Hind margin of carapace without bony nodules; nine or more
plastral callosities............................................. Cyclanorbis
3’ Hind margin of carapace containing bony nodules; seven plas
tral callosities ...................................................... Lissemys
4 Bony arcade behind eye broader than the diameter of the orbit
....................................................................................... 5
4’ Bony arcade behind eye narrower than the diameter of the orbit
...............................................................................Trionyx
5 Head short, broad, and roundish; distance from nasal opening
to orbit about equal to diameter of orbit; no dark longitudi
nal lines on the head ..........................................Pelochelys
5’ Head long and narrow; snout extremely short; distance from
nasal opening to front of orbit much less than the diameter of
the orbit; dark longitudinal lines on the head............ Chitra
17
www.Ebook777.com
Chapter 2
TURTLE STRUCTURE AND FUNCTION
The character that makes the Testudines one of the most
clearly defined of all vertebrate orders is the shell, which, though
superficially paralleled by certain other vertebrates (such as the
armadillos and certain armored fish) is in fact unique among ver
tebrates in its degree of involvement with, and modification of,
such important skeletal elements as the ribs, the vertebrae, and
the shoulder girdle. The primary function of the turtle shell is, of
course, a protective one, and the relative invulnerability of adult
turtles may be an important reason for their present survival in
such a wide variety of ecological niches. However, it is also worth
considering that the possession of a shell, with its humped and
rounded shape, makes a turtle many times heavier and bulkier
than another reptile of comparable body length. To take an ex
treme example, a tree snake with a body length of three feet may
weigh only a few ounces, while a tortoise of that length might
weigh over three hundred pounds. Obviously much of this extra
weight and volume is devoted to supportive and muscular struc
tures, but nevertheless a turtle does have a much greater capacity
for storing food and water than another reptile of comparable
length. The voluminous body also enables the female turtle to pro
duce a large volume of eggs, and some of the larger turtles (for
example the sea turtles and the freshwater sideneck Podocnemis
expansa) lay larger clutches than any other amniote vertebrates.
The small clutches of present-day giant tortoises are an adaptation
to life on simple oceanic island ecosystems, where it is undesirable
to produce too many young. The capacious turtle body is also very
suitable for housing the bulky intestines necessary for digestion of
vegetation. A very large proportion of turtles are, in fact, partially
or completely herbivorous, while there are no vegetarians among
the snakes and crocodilians and only a few among lizards.
The structural details of the turtle shell show considerable
variation throughout the order, but the pattern typical of emydid
and testudinid turtles (which together constitute well over half of
the living species) may be taken as a convenient starting point for
a description of the various elements.
The turtle shell is composed of a convex dorsal part, the cara
pace, and an almost flat ventral part, the plastron; the carapace
18
and plastron meet at a ‘bridge’ on each side. Both carapace and

plastron consist of a superficial layer of homy material, divided
into large plates known as scutes which overlie a layer of inter
locking bones. The word ‘lamina’ is often used instead of ‘scute,’
but since ‘lamina’ has a rather precise meaning in osteology it is
preferable not to use it in this different sense. Scutes are also
sometimes called plates, shields, or corneoscutes.
OSTEOLOGY OF THE CARAPACE

The carapace invariably includes a large, polygonal, anterior
median element, the proneural or nuchal bone, which is present in
all turtles and is an important site of attachment for the neck mus
cles. Behind the proneural is a series of smaller bones, the neurals.
These are almost always hexagonal in shape in emydid turtles, but
are modified into alternating square and octagonal elements, at
least toward the middle of the carapace, in most tortoises. Some
semi-terrestrial emydids (Terrapene, Rhinoclemys) may also
have at least one octagonal neural bone. The neural bones, typical
ly about nine in number, incorporate the neural arches of the trunk
vertebrae. The median rear element is known as the pygal bone,
and those bones between the pygal and the posteriormost neural
bone (defined as the last median bone attached to a vertebra) are
known as suprapygals. In many tortoises two suprapygals are
present, the anterior one with two posterolaterally directed rami
which enclose the oval or semicircular posterior one.
The neural bones are flanked by two series of narrow, curved
bones known as pleurals. Each pleural develops around a rib, and
the total number is very constant at eight on each side. The anteri
or pleural on each side Is pentagonal in shape and larger than
those behind it. In most tortoises the pleurals have alternately
wide and narrow proximal ends, corresponding with alternately
octagonal and square neurals. A pleural wide at its proximal end is
narrow at its distal end, and vice versa. In hatchling turtles, well-
defined ribs are present but pleural bones are virtually absent. As
the turtle grows, the gaps between the proximal ends of the ribs
become bridged by bone, and gradually these bony connections be
come increasingly complete toward the distal (or lateral) ends of
the ribs. When viewed from within, it can be seen that the head of
each rib articulates with a vertebra. In certain turtles with long
and highly extensible necks, such as Deirochelys, Chelydra, and
Chelus, the median part of each rib loops away from the pleural
bone in which its distal part is embedded; the channels thus de
fined on each side of the vertebral column are, in life, occupied by
the neck extensor muscles.
The distal rib-ends do not curve round toward each other as in
other vertebrates, but they may project a little beyond the edges of
19
www.Ebook777.com
Carapace (above) and plastron (facing page) of the tortoise Gopherus
polyphemus. Art courtesy Patrick Burchfield.
20
21
www.Ebook777.com
the pleural bones. The pointed rib tips are inserted in pits in the so-
called peripheral bones, which form a continuous series of about
eleven on each side, the anterior pair being in contact with the pro-
neural bone and the posterior pair with the pygal bone.
The turtle carapace thus forms a strong dome constructed on
the soundest engineering principles. However, such a dome will
only be resistant to vertical crushing if the sides are rigidly an
chored so they cannot spread apart. This is one of the functions of
the belly shell or plastron, the other functions being the obvious
one of providing protection for the underside of the animal, and
also the anterior and posterior parts of the plastron provide a rigid
point of attachment for numerous neck and limb muscles.
OSTEOLOGY OF THE PLASTRON

The plastron is composed of four paired elements, known
(from front to back) as the epiplastra, hyoplastra, hypoplastra,
and xiphiplastra (singular plastron in each case). A diamond
shaped median element, the entoplastron, is located at the junc
tion of the epiplastra and hyoplastra. The entoplastron is thought
to be derived from the interclavicle bone of other reptiles,, the epi
plastra from the clavicles, and the other plastral bones from modi
fied abdominal ribs, or gastralia. Even in baby turtles the front
and back parts of the plastron, important for muscle attachment,
tend to be well ossified, but at birth there is a large fontanelle be
tween the hyo- and hypoplastra which gradually closes with
growth. The plastron connects with the carapace by means of
sutures between the middle peripheral bones (which have an ap
propriate V-shaped cross section) and the hyo- and hypoplastra.
These latter bones also throw up strong buttresses, each of which
connects both with one of the peripheral bones and with either the
first, or approximately the sixth, pleural bone on each side.
VARIANT SHELL STRUCTURES

Although the above description applies to the majority of
living turtle species, exceptions are quite plentiful. In mud turtles
(Kinosternidae), some of the neural bones are eliminated, so that
the rearmost (and sometimes the frontmost) pairs of pleurals
meet on the midline. This condition also obtains in Terrapene and
the trionychids. In certain chelid genera (Platemys, some Phryn-
ops, Chelodina, Emydura, Pseudemydura, and Elseya) the neur-
als are completely lost, and each pair of pleurals meets at a
median suture. In Lepidochelys, on the other hand, the neural
bones may be considerably increased in number by both longitudi
nal and transverse division of individual elements.
22
All living turtle species normally have eight pairs of pleural

bones, though in some American Trionyx species the last two
pleurals on each side may fuse. The constancy of the pleural count
Is probably associated with the fact that each pleural is closely as
sociated with, and grows around, a rib, and ribs in turtles appear
to be much less variable numerically than shell elements. How
ever, the relative sizes of the pleural bones are variable. For ex
ample, in certain chelid turtles with very long necks (Chelodina,
Hydromedusa), the neck cannot be withdrawn posterior to the
level of the first fixed vertebra because of the lateral plane of re
traction, so the head and neck can only be protected by a forward
extension of the front part of the carapace. Slice the first trunk
vertebra is fixed by its associated ribs at the le Tel of the first pair
of pleurals, the forward extension of the carapace has to be pro
duced by a front-to-rear elongation of the front half of the first
pleurals, as well as of the nuchal bone and the first few periphe
rals.
In the soft-shelled turtles, the peripheral bones are completely
missing, and the free rib-ends are simply embedded in the cartila
ginous shell-flap. However, the bony part of the shell of these tur
tles is composed of very thick bone and is so flat that it does not re
quire any further connection between the two sides. In certain
other genera (such as Claudius and Platystemon) peripheral
bones are present, but do not have any sutural connection with the
plastron. In both these genera the carapace is not only very thin,
but also strongly depressed, so one can only assume that these tur
tles have specialized habitats or ways of life that do not expose
them to the same predators as other turtles. In the American box
turtles (Terrapene), the plastron is composed of two moveable
lobes with a hinge between the hyo- and hypoplastron. A rigid
bridge is thus impossible, but the shell derives considerable resis
tance to crushing both by having a steeply domed shape with a
stout or flared rim and by developing complete fusion between all
elements of the carapace, and of each lobe of the plastron, when
adult size is reached. In most other turtles, the various bony shell
elements remain distinct even in large adults.
In certain aggressive and predatory turtles (such as Chelydra
and Staurotypus), ventral armor becomes unimportant and the
plastron becomes reduced to a cruciform shape (which of course
still retains the functions of bracing the sides of the carapace and
of providing anterior and posterior muscle attachments). In two
closely related genera (Kinostemon and Stemotherus), the ento-
plastron disappears and both front and rear parts of the plastron
become attached by flexible hinges to the rigid middle part of the
plastron (composed of the hyo- and hypoplastra). In the supposed
ly related genus Claudius, however, an entoplastron is present and
23
www.Ebook777.com
Carapace (facing page) and plastron (above) of the soft shell Trionyx ferox.
Parts of plastron: A - preplastra; B - fused epiplastra; C - fused hyo- and
hypoplastra; D - xiphiplastron. Art courtesy Patrick Burchfield.
hinges are absent, and the paired bony elements of the plastron
are reduced to three on each side, the middle pair taking the place
of both hyo- and hypoplastra.
In many long-extinct turtle genera one or two extra pairs of
bones, known as mesoplastra, were intercalated between the hyo-
and hypoplastra. Among living turtles only the three genera of the
Pelomedusidae retain one pair of mesoplastra. In Pelusios the
mesoplastra are quite large and still have a median connection,
but in Podocnemis and Pelomedusa they are small and widely-
spaced. Mesoplastra may be present as a rare variant condition in
certain cryptodires (Lepidochelys, Malacochersus).
The entoplastron is absent in soft-shelled turtles (Trionychi-
dae), but the epiplastra are completely fused and neomorphic
structures known as ‘preplastra’ are present. The bony plastron of
trionychids is highly reduced; the front part of the plastron is flexi
ble and can be raised when the head is retracted.
25
www.Ebook777.com
Bony shell of the aberrant African pancake tortoise, Malacochersus tornieri,
showing reduction of bony area. Redrawn from Procter, 1922.
SCUTE TERMINOLOGY
In most turtles the bony shell is covered with a superficial
layer of horny scutes. These vary greatly in thickness, from paper-
thinness in Dermatemys to nearly a quarter of an inch thick in
Eretmochelys. Typically there is a median series, of five large
scutes known as vertebrals or centrals, flanked by two rows each
composed of four costals, or laterals. The scutes around the edge
are known as marginals, with a small anterior nuchal and a large,
sometimes longitudinally divided, posterior supracaudal. It has
been argued that the term ‘supracaudal’ properly refers only to
structures on top of the tail itself (such as the supracaudal scales
of snapping turtles), and that the substitute term ‘postcentral’
should be used. But rigid application of this rule would require
changes in the names of most of the plastral scutes, which would
be most unfortunate since current plastral scute nomenclature is
distinguished by its stability and universal acceptance.
The bones near the middle of the carapace are smaller than
the overlying scutes, and one central scute may cover parts of up
Generalized scutes of the carapace (left) and plastron (right). Carapace: N

nuchal; C - centrals; Co - costals; M - marginals; S - supracaudals. Plastron: I
Intergular; G - gular; H - humeral; P - pectoral; Ab - abdominal; F - femoral; An
Anal.
www.Ebook777.com 27
to nine or ten bones. However there is usually a one-to-one corres
pondence between the peripheral bones and the marginal scutes,
each marginal covering half of each of two peripherals. The scutes
of the plastron are arranged In six pairs, known respectively as the
gulars, humerals, pectorals, abdominals, femorals, and anals. The
names are derived from the parts of the body that lie beneath. A
small scute in each armpit is called the axillary, and a correspond
ing one in front of each thigh as the inguinal.
VARIANT SCUTE CONDITIONS

Variations on the theme outlined above are rather plentiful. In
Caretta five pairs of costals are present, and in Lepidochelys there
are at least five, and sometimes as many as nine or ten. In Noto-
chelys six (or seven) centrals are present, and in Lepidochelys oli-
vacea there may be anywhere from five to nine. In Hydromedusa
the nuchal scute is excluded from the front margin of the carapace
by the anterior marginals, and it is considerably enlarged, giving
it the appearance of an extra central. In many tortoises and side
necked turtles the nuchal scute is missing. In sidenecked turtles
(Pleurodira) and sea turtles (Cheloniidae) a median anterior plas-
tral scute, known as an intergular, is present. In the Cheloniidae,
Platystemidae, Dermatemyidae, and Chelydridae, a row of scutes
is present between the axillary and inguinal on each side, separa
ting the main plastral scutes from the lower part of the marginals;
these scutes are known as inframarginals. In Macroclemys a row
of three or four scutes is intercalated between the middle margi
nals and the costals on each side; these are known as supramargi-
nals. In Kinostemon only a single gular is present, and in Stauro-
typus and Claudius the plastral scutes are reduced to four pairs.
Although all turtle species except Lepidochelys olivacea have
a ‘standard’ carapace scute conformation, individual variants,
usually but not always asymmetrical, are quite plentiful, though
the frequency of variation is very different from one species to
another. Certain definite types of irregularity are recognizable,
one of the most distinctive being doubling (or near doubling) of the
number of vertebral scutes, which are arranged in alternating,
staggered pattern. In this particular deformity the intercostal
seams alternate, there usually being one costal more on one side
than the other.
Much scholarly sweat has been expended on attempts to homo-
logize shell scutes of ‘deviant’ turtles with scutes of the ‘standard’
turtle shell. Such efforts, however, are probably pointless. One can
speak of homology between bones or structures of different verte
brates (such as the arm of man and the wing of a bat) because it is
possible to demonstrate the gradual divergence of both of these
parts from a common origin, by both ontogenetic studies (of em
28
bryos) and phylogenetic studies (of fossils). However, there is no

scute pattern standard for all turtle embryos which gradually
diverges in different turtles Into the various scute patterns of the
adults. Instead, the scales are formed in their final adult arrange
ment at a fairly early embryonic stage, each scute being delimited
by a wrinkle in the still-soft carapace epidermis. Thus, if a turtle
of a species which normally has six pairs of plastral scutes produ
ces an offspring with only four pairs, there is no scientific meaning
to any argument as to whether the gular and humeral scutes have
been eliminated, or the gulars and anals, or the pectorals and
anals.
It is in fact remarkable how common the ‘standard’ (i.e. testu-
dlnld or emydid) scute pattern is among turtles. It is hard to see
any particular virtue in having five central scutes and four pairs of
lateral scutes; yet this arrangement is found even in turtles which
have had no common ancestor since the Jurassic. On the other
hand, individual turtles with highly abnormal scute patterns do not
usually seem to be at any discernible disadvantage.
Different parts of the turtle shell are not equally liable to ab
normality. Both the bones and the scutes of the plastron are re
markably stable, as are the peripheral, pleural, and proneural, or
nuchal, bones. The neural and suprapygal bones, however, may
vary considerably both in number and in shape even within a spe
cies. Extra vertebral scutes are moderately common, as are extra
or reduced costal scutes; both are more common than abnormal
numbers of marginals.
In two turtle families (Trionychidae and Dermochelyidae) the
shell scutes have been eliminated, while in a third family (Caret-
tochelyidae) they are only indicated by vestigial vertebrals in
some hatchlings. In all three families the carapace and plastron
are covered with soft skin.
EPITHECAL OSSIFICATIONS
Epithecal ossifications, or superficial bony structures, are
found in the shells of a few turtle species. The leatherback turtle
(Dermochelys) has a continuous layer of small, irregularly
shaped mosaic bones between the ribs and the surface skin. Some
adult soft-shelled turtles (Trionychidae) develop superficial
heavily sculptured bony callosities on both the carapace and the
plastron, and old matamata turtles (Chelus) may develop epithe
cal bony caps on some of the carapace tubercles.
SHELL GROWTH AND PIGMENTATION

Growth of the turtle shell is much more complicated than that
of shelled invertebrates, in which the shell generally enlarges by
www.Ebook777.com 29
simple deposition of new calcified material over the inside surface
and at the exposed edges of the shell. The bony shell of a turtle is
quite well-supplied with blood vessels, and new bone is added
around the edges of each of the individual bones of the shell. In
hatchling turtles the bones in many cases are not in contact with
each other, but they grow outward toward each other and, when
they eventually meet, form sutural connections. Growth then con
tinues at the sutures, though increasingly slowly as adult size is
reached, and in very old turtles (or in quite young adult American
box turtles) the shell sutures may ossify completely, whereupon
growth ceases. In some turtles certain of the shell sutures may be
come secondarily invaded by soft tissue. The breakdown and
resultant hinges in carapace and plastron have several functions,
among which are further protection of the extremities and facilita
tion of respiration and oviposition.
The scutes grow by gradual deposition of new horny material
over their entire undersurface by a layer of Malpighian or horn-
secreting cells on the surface of the bone. These cells propagate
laterally as the shell grows, so the bony shell is always uniformly
covered by them. In many turtles—especially those in climates
with a cold winter—growth proceeds in cycles, and a wrinkle is
formed around the edge of the scute as growth ceases at the end of
the season. Thus the age of some turtles may be calculated by
counting the number of rings around each scute—at least when the
turtle is young and the growth rings are not too crowded. For tur
tles under the artificial conditions of captivity, however, many
other factors than the onset of winter will slow or stop growth, and
this method of calculating the age can no longer be used. In theory
it would seem that the central part of the scute would become
much thicker than the edges as growth proceeds. However, in
many aquatic turtles the outer layers of the scutes peel off at quite
frequent intervals, so that the scutes remain thin and devoid of
growth rings. Also, in tortoises the central parts of the scutes be
come worn down by mechanical abrasion, and it also seems that
new keratin is laid down more abundantly at the edges of the
scutes than near the middle.
One scute can cover parts of several bones; a vertebral scute,
for example, may cover parts of four neural bones, three left
pleurals, and three right pleurals. Consequently much
sliding-about of scutes over bones occurs as a turtle grows. How
ever, the original or juvenile part of a scute (the areola) tends to
remain attached to that part of the bony shell which was originally
below it. The result is that in an adult turtle the areolae are fre
quently far from centrally located on the scutes, and the growth
rings may be tightly packed on one or two sides of a scute and
widely spaced on the other sides. Scute borders, or seams, usually
30
Cirowth of the common box turtle, Terrapene Carolina, natural size. Top to
bottom: one month, one year, two years. Photos by R.J. Church.
31
www.Ebook777.com
remain clearly defined even in old turtles, though in large speci
mens of Batagur or Dermatemys they may become indistinct.
The scutes are far from a passive covering for the bones be
neath. The scute borders usually produce strong furrows, known
as sulci (singular sulcus) in the bone below, which are often much
more obvious on a complete bony shell than the actual bony su
tures. In some turtles (such as Malaclemys) even the growth rings
of the scutes may produce strong furrows in the underlying bone.
Also, under captive conditions tortoises sometimes grow in an ab
normal fashion in which each of the large carapace scutes forms a
markedly conical shape which is reflected in the shape of the
underlying bones. Conical dorsal scutes are present to a less mark
ed degree in many wild tortoises.
Some of the common patterns of shell pigmentation in turtles
are direct results of the manner of scute growth. For example, in
rapidly growing turtles, melanin production may not keep pace
with keratin production, and the individual scutes will be dark with
white or horn-colored edges (as in the carapaces of young Galapa
gos tortoises or the plastra of certain Rhinoclemys species). Ano
ther thing that can happen is that the melanin may be distributed
in clumps rather than uniformly in the scute at birth; as the scute
grows each melanin clump propagates itself outwards, forming a
dark radial streak, while from points where no melanin was pres
ent initially a light radial streak is formed. The effect of this is to
produce a beautiful starred pattern, exemplified by such tortoises
as Geochelone radiata and G. elegans. In Geochelone carbonaria,
on the other hand, the carapace is fairly light in color at birth, and
melanism is only produced in the postnatal growth of each scute.
Consequently in the adult the carapace scutes are uniformly black
except for a single, circumscribed, yellow spot on each.
In many turtles (such as Pseudemys, Deirochelys, and More-
nia), more complex patterns may be present on the carapace
scutes, such as non-radial streaks, eccentric ocelli, or irregular
spots. These pigment patterns are sometimes evident on the shell
bones as well, though they usually terminate abruptly at the edge
of a scute. The formation and growth of such patterns are incom
pletely understood; they usually break down in large adults, in
contrast with the permanent shell patterns of most starred tor
toises.
SKULL STRUCTURE
In the earliest fossil turtles, and in the modern families Chelo-
niidae, Dermochelidae, and Platystemidae and the sideneck spe
cies Peltocephalus tracaxa and Pseudemydura umbrina, the
head is non-retractile and the skull has a complete bony roof.
32
Viewed from behind, the skulls of these turtles show two large dor
sal cavities, bounded by the supraoccipital, parietal, squamosal,
and paroccipital bones. These cavities in life are occupied by the
powerful jaw muscles. The much smaller brain, or cranial, cavity
is situated in a separate bony chamber below these muscles. The
occipital condyle—the point of articulation with the anteriormost
cervical vertebra, or atlas bone—is located below the opening for
the spinal cord, or foramen magnum, and is composed of a single,
cartilage-surfaced knob, as in all reptiles. This knob, however, is
made up of parts of three separate bones, the basioccipital and two
exoccipitals. The roof of the skull is composed of paired prefront-
als, frontals, postfrontals, and parietals, all very stout bones in
Dermochelys, but less so in some other sea turtles (e.g. Lepido-
chelys olivacea and Caretta caretta). The jaw margin is composed
of the paired premaxilla and maxilla bones. The hind parts of the
skull and ear region are made up by the jugal, squamosal, and
quadrate bones, while sound is transmitted from the external
membrane to the inner ear by the slender columella bones. The
Skull of the common snapping turtle, Chelydra serpentina , illustrating the

principal bones. Redrawn from Ashley, 1962.
33
www.Ebook777.com
palatal region is composed of a single vomer, paired palatines and
pterygoids, and a single basisphenoid. The lower jaw is composed
of a single U-shaped dentary, with paired coronoid, articular, sple-
nial, and surangular bones near the point of articulation with the
skull. The hyoid structure in the base of the tongue is often large,
with two pairs of well-ossified rami.
It is probable that the complete skull roof of modem sea tur
tles is a primitive hold-over from their cotylosaurian ancestors;
certainly there is no evidence that the skull roof was lost and rapid
ly re-formed between the Triassic ancestral turtles and the Creta
ceous cheloniids. The thick, fleshy neck of sea turtles, necessary
for effective streamlining, has the result that the head cannot be
retracted into the shell, and the skull roof was presumably retain
ed to make the head less vulnerable to attack. However, Platyster-
non is a comparatively modem genus, specialized for climbing
trees and steep places. The head in this form is very large, so that
the body is counterbalanced when climbing, and this large head,
not being retractile into the very depressed shell, again requires
an intact skull roof for protection, though in this case it may well
be a secondary re-development.
In all turtles apart from the ten species included in the fami
lies and genera mentioned, the head is more or less retractile and
the skull roof is emarginated to allow the jaw muscles to bulge out
when the jaw is forcibly closed; in the roofed-over turtles these
muscles can bulge to a limited extent by means of a complex fluid-
displacement arrangement. The skull emargination usually pro
ceeds from the back, at the rear of the parietal bones, and may
proceed so far that only a narrow temporal arch remains; in some
emydid genera (e.g. Heosemys) even the arch disappears. In one
family, the Chelidae, this emargination starts from below, and
proceeds upwards and backwards; in one chelid genus (Chelo-
dina) the arch disappears completely.
Gaffney (1975) recently summarized a series of interesting
papers by Schumacher (1954, 1955a, 1955b, 1956) describing the
unique jaw-closing system of turtles. Jaws in most vertebrates are
closed by contraction of a bunch of temporal muscles centered
around an adductor tendon known as the Bodenaponeurosis.
Usually the bodenaponeurosis runs directly from its dorsal
attachments in the temporal area to the coronoid process on the
jaw, but in turtles the expanded otic chamber mae such a straight
connection impossible. Accordingly, the cryptodiran and
pleurodiran turtles have independently evolved a trochlear, or
“pulley”, mechanism for transmitting the jaw-closing tension
around the otic chamber. In cryptodires, the jaw muscles pass
over the smooth, saddle-shaped, antero-dorsal surface of the
chamber, the area known as the processus trochlearis oticum.
34
That part of the bodenaponeurosis that passes over the otic

chamber contains an embedded cartilaginous nodule known as the
cartilago transiliens. This small cartilage is meniscus-shaped,
with the concavity fitting the surface of the trochlear. In certain
tortoises (e.g. Gopherus) this element becomes bony rather than
cartilaginous, in which case it is known as the os transiliens. In
pleurodires, the trochlear is formed in an entirely different way,
by means of lateral processes from the pterygoid, each known as a
processus trochlearis pterygoidei. A cartilago transiliens is
present in pleurodires, but instead of forming a synovial capsule
the “joint” is formed by an infolding of the mucous membrane of
the mouth.
Topological deformations of the ‘normal’ shape turtle skull are
legion, but actual alterations in the numbers of bones are rare. In
the Chelidae, however, only a single premaxilla is present, and the
two sides of the lower jaw may not be fused.
The alveolar, or biting, surfaces of the jaw vary greatly ac
cording to diet. In many mollusc-eating species an extensive
secondary palate is formed, and both upper and lower jaws are
equipped with heavy crushing plates. In many herbivorous species
(terrestrial, freshwater, and marine), the alveolar surfaces have
tooth-like serrations, while in predatory and scavenging species
the jaw surfaces may be exceedingly sharp. True teeth are un
known in turtles with the exception of a few palatal teeth in the
earliest known fossil genera (Proganochelys and Triassochelys),
and also palatal teeth have been recorded from certain embryonic
soft-shells.
CERVICAL AND DORSAL VERTEBRAE

In the earliest turtles, seven cervical vertebrae and eleven
trunk vertebrae were present, but this was soon altered to eight
cervicals and ten trunk vertebrae, an arrangement that has re
mained stable and universal to the present day. The actual verte
bral structure is very interesting and is perhaps the most special
ized, and certainly the most differentiated, among vertebrates.
Normally vertebrae serve several important functions—those of
protection of the spinal cord and dorsal arteries, of providing
muscle attachments, and of constituting the basic elements which
define the shape and proportions of the animal. However, the dor
sal vertebrae of turtles are relieved of all of these functions, and it
is not surprising to find that they have degenerated considerably.
Only the anterior end of the first dorsal vertebra retains some
complexity, since it requires an articular connection with the
eighth cervical vertebra. The first dorsal vertebra lies under the
35
www.Ebook777.com
Cervical vertebrae of: Hieremys annandalei (emydid) (left); Lepidochelys kem-
pi (sea turtle) (center); and Chelus fimbriatus (side-neck) (right). Redrawn from
Williams, 1950.
36
nuchal bone, and the two have strong cartilaginous (but not bony)
connections. This vertebra is fixed in place posteriorly by the dor
sal ends of the first pair of ribs, and anteriorly by the so-called
“costiform processes” of the first pleural bones. Anteriorly there
is a cotyle, or concavity, for reception of the posterior convexity,
or condyle, of the eighth cervical. The first dorsal vertebra also
has prezygapophyses which articulate with the postzygapophyses
of the eighth cervical. These are widely separated in cryptodires,
but close together near the midline in sidenecks (to facilitate late
ral flexion of the neck).
The other nine dorsal vertebrae are composed of two parts,
with a horizontal plane of fusion. The more dorsal part, the neural
arch, is simply a longitudinally grooved projection from the vis
ceral surface of a neural bone, with blunt lateral processes at the
midline which articulate with a rib. The ventral part, or centrum,
is a dorsally grooved elongate rod-like bone, with expanded ends
whose flat surfaces are separated from their neighbors by fixed
cartilaginous discs. The centra of the trunk vertebrae are ar
ranged in staggered fashion with respect to the neural arches, so
that each centrum covers the posterior half of one neural arch, and
the anterior half of another. Because of the staggering, the rib-
ends connect with the junctions of the centra of the vertebrae,
rather than with the mid-point of each. The dorsal vertebrae and
dorsal rib-ends are fairly substantial in many turtles, but in some
(for example many tortoises) the dorsal vertebrae are extremely
lightly constructed, and the rib-ends may become so thin that they
almost disappear. In adult Malacochersus the ribs disappear com
pletely. In some chelid turtles neural bones are absent and conse
quently the trunk vertebrae are represented only by centra; the
dorsal part of the spinal cord is housed within a groove at the medi
an junction of the pleural bones.
To compensate for the rigidity of the body, the neck of the tur
tle is relatively very long and highly flexible. The classical type of
reptilian locomotion involves throwing the body into horizontal sig
moid curves, and this movement is retained (or re-formed) in the
cervical region of sideneck turtles (Pleurodira). In these turtles
lateral flexion is made possible at each of the cervical articula
tions either by a ball-and-socket joint or by joints in the form of
vertical semi-cylinders fitting into vertical grooves in their neigh
bors. In the Pelomedusidae all the cervical vertebrae are normal
ly procoelous (i.e., with a concavity in front and a convexity be
hind) except for the axis, or second cervical, which is biconvex,
and the atlas, or first cervical, which is amphicoelous (concave at
both ends), as it is in virtually all turtles. In some species saddle
surfaced joints may also be present. In the Chelidae, however, the
second to fourth cervicals are opisthocoelous (i.e. with a convexity
37
www.Ebook777.com
in front and a concavity behind), the fifth is biconvex, the sixth
procoelous, the seventh amphicoelous, and the eighth again bicon
vex. The otherwise somewhat loose pleurodiran cervical structure
is stabilized by well-defined pre- and postzygapophyses, arranged
so as not to restrict lateral flexure. There are well-defined lateral
processes throughout, to provide attachment for the neck-flexing
muscles (the Musculi testo-cervicalis lateralis of the longissimus
system).
The sagittal flexure of the neck of cryptodiran turtles has
necessitated radically new types of cervical connections. The most
striking feature of the cryptodire neck is the doubling of the proxi
mal articular condyles to restrict lateral bending of the proximal
part of the neck. These double condyles are known as ginglymes,
and the condition as ginglymoidy. In one species (Carettochelys
insculpta) there is a double (procoelous) joint between the first
vertebral and the eighth cervical, but in all other cryptodires there
is a single procoelous joint at this point (except in the Trionychi-
dae, where this connection is invaded by ligamentous material and
there is no bony joint; in the retracted neck of a trionychid these
vertebrae lie belly-to-belly).
Almost all cryptodires (except for occasional snapping turtles
and sea turtles) have a double joint at the anterior end of the
eighth cervical. These articulations are convex-forward in Platy-
stemon, all emydids, and all testudinids (except for some speci
mens of Pyxis arachnoides). Carettochelys and trionychids are
also convex-forward at this point, but sea turtles, snapping turtles,
Dermatemys, and kinostemids have a concave-forward anterior
eighth cervical articulation. The anterior end of the seventh cervi
cal Is doubly concave in most cryptodires, but doubly convex in
Carettochelys and many trionychids, and frequently flat in sea
turtles. In the Carettochelidae and Trionychidae all the other cer
vical vertebrae are single-ended opisthocoelous (except for the
amphicoelous atlas). In other turtles ginglymoidy may still be pre
sent at the anterior end of the sixth cervical (some emydids, testu
dinids, and Platystemon), and anterior opisthocoely is reached by
means of a biconvex vertebra at position 4 in sea turtles, snapping
turtles, Platystemon, some emydids and testudinids, at position 3
in some kinostemids, emydids and testudinids, and at position 2 in
Pyxis, Dermatemys, and some Kinostemon. The atlas vertebra is
amphicoelous in all turtles except for a minority of specimens of
Pyxis.
Well-defined zygapophyses are present throughout the crypto
dire cervical vertebrae. The prezygapophyses have convex sur
faces and face upward, while the postzygapophyses have concave
surfaces and face downward; this arrangement permits free sli
ding of post over prezygapophysis as the neck is bent in the normal
38
sagittal plane. Each joint does not contribute equally to the flex
ure; kinesis is concentrated at the joints at each end of the eighth
cervical, and in testudinids and emydids there is an anterior cen
ter of movement, near the middle of the neck, posterior to the bi
convex centrum. Sea turtles theoretically also have this anterior
(enter of movement, but the cervical vertebrae are so massive
and allow so little flexure of the neck that it is scarcely functional.
In many cryptodires the cervical vertebrae retain a horizontal
plane of cleavage, or suture, between the centrum and the neural
arch, which may persist even in adults. The rather specialized at
las, unlike that of pleurodires, often remains incompletely ossified
throughout life, the centrum, intercentrum, and two halves of the
neural arch being connected only by cartilage and separating on
maceration. Vestigial ribs may be present at certain cervical
Joints, remnants of the cervical ribs of the earliest turtles such as
Proganochelys.
SACRAL VERTEBRAE
In cryptodires, the ten trunk vertebrae are followed by two
sacral vertebrae. Both connect with sacral ribs whose broadened
ends meet the iliac crests of the pelvis. The sacral ribs are very
slender in the Testudinidae, and the similarly slender ribs attach
ed to the ninth and tenth trunk vertebrae are directed posteriorly
so that they also reach the iliac area.
In pleurodires the pelvis is firmly fused to both carapace and
plastron, and the distinction between dorsal, sacral, and caudal
vertebrae is less distinct. However the ribs of the seventeenth,
eighteenth, and nineteenth (and sometimes the sixteenth and
twentieth) vertebrae are fused to the centra and reach the ilia.
CAUDAL VERTEBRAE
The number of caudal vertebrae is variable, but usually lies
between 25 and 30. The turtle tail is capable of muscular move
ment throughout its length, and the caudal vertebrae, except to
ward the very tip, are well-defined and in no way degenerate. The
general form is similar to that of a cervical vertebra, with strong
pre- and postzygapophyses, though never with ginglymes. Lateral
rib-like processes are well-defined on caudal vertebrae 4, 5, and 6,
and sometimes others. The centra are procoelous, except for the
distal parts of the tail of the very long-tailed forms (Chelydridae
and Platystemidae) where they are opisthocoelous, with a single
biconvex centrum at the changeover. Haemal arches are present
from about the eighth caudal onwards, and separate chevron
bones are present in the tails of snapping turtles. Open sutures be
tween the caudal centra and neural arches may persist to maturi-
www.Ebook777.com 39
ty, but usually the two parts become fused, the fusion commencing
with the more distal vertebrae. In some tortoises and kinostemids
the tail tip is armed with a spur composed of a conical bone en
cased in horn. This spur, which curves downward somewhat at the
tip, assists the male in holding his grip on the female during copu
lation. In some recently extinct Mascarene tortoises the bony tail
spur reached a length of up to six inches.
LIMB GIRDLES
The limb girdles of turtles are housed within the rib cage, a
condition unique among vertebrates, and one which requires a
major feat of embryonic legerdemain for its inception. The pur
pose of the girdles is to provide a strong socket to house the head of
the humerus or femur, and to brace this socket in such a way as to
resist the upward, inward force of the thrusting, weight-bearing
limbs. Consequently both girdles have a strong upward process
which rests against the carapace, and mesially-directed processes
which meet, or almost meet, on the midline.
The pectoral girdle is composed of two pairs of bones; the
large L-shaped scapula consists of a long vertical ramus which
rests on a cartilaginous pad beside the first dorsal vertebra, an
acromial process which is directed mesially and slightly anterior
ly to meet its fellows at the midline, and a glenoid fossa, or socket,
which holds the head of the humerus. The other pectoral bone is
the coracoid which is an elongate, spatulate bone directed inward
ly and posteriorly from the glenoid fossa. It is connected by its dis
tal end to the acromial process of the scapula by a ligament. The
coracoids do not meet mesially, so each half of the pectoral girdle
is free to rotate in an approximately vertical plane around an axis
extending from the junction of the scapula with the carapace and
the medial junction of the acromial process.
The pelvis is composed of three pairs of bones—the ilia, ischia,
and pubes. The six bones comprise a rigid unit which has
relatively little free movement within the shell (no free movement
in pleurodires). The pelvis is basically U-shaped, the two sides of
the U being made up of ilia which curve slightly toward each other
dorsally. The upper end of each ilium rests on a bony pedicle ex
tending downward from the inside surface of the eighth pleural
bones. The lower part of the U is expanded from front to rear, the
anterior part being composed of the paired pubes and the posterior
part of the paired ischia. The three bones on each side meet at the
acetabulum, a large socket which articulates with the head of the
femur. On either side of the median fusion of the ischia and pubes
is a large round hole, the obturator foramen. The pubes extend for
ward mesially, ending in a cartilaginous projection known as the
epipubic cartilage. Antero-laterally each pubis terminates in a
40
Skeleton of an emydid turtle, Emys orbicularis, from below, with the plastro
removed. Redrawn from Bojanus, 1819.
41
www.Ebook777.com
strong pectineal process, for attachment of the attrahens pelvim
and retrahens pelvim muscles. The Ischia posteriorly form a
strong attachment for the dilatator cloacae muscles.
In the sidenecked turtles (Pleurodira) each of the six pelvic
bones is firmly fused to the shell (the ilia to the carapace and the
ischia and pubes to the plastron). The need for median fusion be
tween the various bones is therefore eliminated, and the pelvis,
once removed from the shell, is seen to be composed of two sepa
rate halves which do not even touch each other mesially. Conse
quently the median separation between the obturator foramina
disappears, and instead of two holes there is a single very wide
one.
LIMB BONES
The proximal parts of turtle limbs are built on the normal
tetrapod pattern of a single proximal shaft (humerus in the fore
limb, femur in the hind limb), and two parallel more distal shafts
(radius and ulna in the forelimb, tibia and fibula in the hind limb).
The bones are angled in an awkward-seeming fashion, with the
humerus or femur extending more or less horizontally rather than
vertically as in mammals. This has the result that weight-bearing
forces, instead of being borne by vertical shafts of bone, are car
ried largely by constant tension in the joints at each end of the
humeri and femurs. These bones are therefore short and massive,
with greatly expanded ends, while the limb muscles too, especially
in large tortoises, are extremely bulky and strong. The humerus
and femur are similar in appearance, though the humerus is often
markedly curved and the femur usually has a larger head. Beside
the ball-shaped head of each bone there are two proximal tuberosi
ties separated by a groove. Distally both humerus and femur are
expanded into a broad roller-type joint which receives the proxi
mal articulations of the radius and ulna, or in the case of the
femur, the tibia and fibula. These more distal bones are not notice
ably curved and are simple shafts with expanded ends. The ulna is
thicker than the radius, and the tibia thicker than the fibula.
FOOT STRUCTURE
The chelonian foot structure is so variable that it was used as
the principal basis for classifying turtles in the early nineteenth
century. However, the foot structure is an adaptive condition,
while fundamental classifications are better based on less adap
tive, less plastic criteria.
Most turtles have pentadactyl limbs, in the typical condition
there being five well-developed, claw-bearing digits in the forefoot
and four well-developed, claw-bearing digits with a reduced, claw
42
less fifth digit in the hindfoot. However, in some tortoises one of the
digits of the forefoot is eliminated (digit 4 in Testudo horsfieldi,
digit 5 in Homopus areolatus). In Pelomedusa five claws are pres
ent on both front and hind limbs, while in the soft-shells (Triony-
chidae) only three claws are present on each foot. Sea turtles
(Cheloniidae) only have one large claw on each foot, though a
small or rudimentary second claw is often present. The leather
back turtle (Dermochelys) is entirely clawless.
In all turtles the distal ends of the radius and ulna present a
linear expansion which articulates with various polygonal bones
known as carpals. The more proximal carpals have individual
names: that which articulates with the radius is called the radi-
ale; that which articulates with the ulna is the ulnare; while the
bone between these two elements is the intermedium. Another
bone, the proximal centrale, is sometimes present between the
intermedium and the radiale, and a small pisiform may be present
outside the ulnare. Two or three medialia are present distal to the
proximal centrale and the intermedium. Distal to this, the indi
vidual digits become distinct, each being composed of a carpal, a
metacarpal, and two or three phalanges. The actual shape of the
digital bones varies enormously, these being highly elongated in
sea turtles and extremely short in tortoises. Most turtles have two
phalanges in the first and fifth digits, and three in each of the other
three, but tortoises have only two in each digit.
In most tortoises the carpal bones show some degree of fusion.
Common patterns of fusion involve amalgamation of the radiale,
proximal centrale, and second medials. In others, the ulnare and
the fifth centrale become fused, and the first three carpals fuse
also. The metacarpals may become fused to the proximal phalang
es, or the medialia may fuse together. In addition, the pisiform is
frequently lost. When this happens the Musculus flexor carpi ul-
naris inserts on a sheet of connective tissue that surrounds the car
pus. Carpal fusion is most advanced in the extinct Geochelone
grandidieri from Madagascar, in which the radiale, proximal cen
trale, medialia 2 and 3, and carpals 1 and 2 are all fused into a sin
gle large bone. Moreover, in all except the fifth digit the metacar
pal is fused to the proximal phalanx.
The hind feet are somewhat simpler in structure than the fore
feet. Typically six tarsals are present, though this is the result of
considerable fusion of more numerous embryonic elements. Each
toe has a single metatarsal; the first and fifth toes have two pha
langes, while the others have three each.
Different tortoises walk on different parts of the front feet. For
example, in Geochelone the foot bones angle forward sharply at
the carpal level, so that the gait is essentially plantigrade, with the
weight being taken by a fleshy cushion between the carpals and the
www.Ebook777.com 43
substrate. Other genera, such as Kinixys, walk on the toes, the
weight being taken by the phalanges. This condition is known as
digitigrade. The burrowing tortoises of the genus Gopherus have a
rather rigid forefoot, with projecting spike-like claws; these tor
toises walk on their claws (unguligrade).
HYOID BONES
The hyoid bones are embedded in the flesh of the tongue and
neck and have no connection with the rest of the skeleton. The
hyoid is made up of a central body and two pairs of postero-lateral-
ly directed horns. The two pairs of horns correspond to the third
and fourth visceral arches of primitive jawless vertebrates; the
jaws correspond to the first arch and the second arch is represent
ed by small, frequently cartilaginous nodules on either side of the
anterior part of the hyoid body, or Basis linguae. The bony nodule
in the tip of the tongue, known as the Os entoglossum, frequently
fuses with the Basis linguae. The hyoid body has a longitudinal
groove in the dorsal side in which the trachea, or windpipe, lies.
There is a hole in the anterior part of the hyoid body known as the
fovea fenestrata, through which the larynx passes.
MUSCULAR SYSTEM
The muscles of turtles have in many cases undergone radical
alteration from the typical vertebrate condition, as one might ex
pect for an animal whose body is encased in a rigid shell. For ex
ample, those muscles which normally produce flexion of the dorsal
spine become completely obsolete in turtles, in which the dorsal
spine is rigid. However, the complex movements of the long and
mobile neck require numerous well-developed muscles, and the
limbs and tail too are highly muscular. The muscles of one turtle
species, Emys orbicularis, have been described and illustrated in
beautiful detail by Ludwig Bojanus in his Anatome Testudinis
Europeae, published in 1819 (reprinted in 1970 by the Society for
the Study of Amphibians and Reptiles), and a few variants from
this typical structure have been described in the technical litera
ture from time to time. However, variations in musculature
throughout the Order Testudines are far from thoroughly studied.
The account given here is based upon Laboratory Anatomy of the
Turtle by Laurence M. Ashley, and refers to a typical emydid tur
tle (Pseudemys).
Within the head itself, the only movements are opening and
closing of the eyes and mouth, and movements of the eyeballs. The
eye is opened and closed by the Musculus levator palpebrae super-
ioris. Focusing of the eye is carried out by the intra-ocular muscles
around the lens, while movement of the eyeball is controlled by the
44
Muscles of an emydid turtle from below, the plastron removed. Redrawn from
Ashley, 1962. See the chart on the following pages for more detailed informa
tion.
45
www.Ebook777.com
www.Ebook777.com
48
www.Ebook777.com
four rectus muscles which have their origins near the center of the
deepest part of the orbit; the superior rectus inserts dorsally on
the eyeball, the inferior rectus ventrally, and the anterior and pos
terior recti in the positions suggested by their names. There are
also two oblique muscles, the superior and inferior, which have
their origins on the interorbital septum. In addition there is the flat
pyramidalis muscle covering the antero-ventral surface of the
eye.
The powerful temporal muscles, which close the jaws, insert
on the coronoid bones and pass postero-dorsally outside the ptery
goids and the cranial cavity, finally grading into supratemporal
fascia which attaches to the occipital process. The jaws are opened
by much weaker muscles, the digastrics, which have their origin
on the ventral side of the tympanic bulla and insert beneath the
posterior ends of the jaws.
Movements of the extremities are complex and are controlled
by large numbers of muscles. These may be divided into those of
the head and neck, of the pectoral girdle and forelimbs, of the pel
vic region and tail, and of the hindlimbs. The names, positions, and
functions of these muscles have been summarized by Laurence
Ashley.
It should be mentioned that in a number of turtle genera the
plastron is hinged and the anterior or posterior parts, or both, can
be raised with very considerable force. The front lobe is raised es
sentially by the action of the pectoralis major in such turtles, and
the hind lobe by the oblique abdominis and transverse abdominis.
In addition to the muscles outlined above, it should be mention
ed that some aquatic turtles, notably Lissemys punctata, have a
sheath of striated muscle, the Muscularis striatum pulmonare,
which envelops the lungs and presumably assists in expiration.
This muscle partially envelops the lungs in Melanochelys, but is
absent in tortoises and in Trionyx. The muscle only covers the
anterior part of the lungs in Podocnemis; in this genus the anterior
portion of the medial part of the muscle sheath arises from the
body of the third thoracic vertebra, passes over and adheres to the
lungs, and has its final insertion on the dorso-lateral part of the
pericardium. The action of a muscle such as this would appear to
be one in which the emptying of the lungs was followed by empty
ing of the auricles, or else one in which the lung pressure is slightly
altered with each heartbeat.
TURTLE RESPIRATION
Respiration presents special problems for turtles, especially
those not equipped with the Muscularis striatum pulmonare, since
the usually rigid shell enclosing the entire visceral mass prevents
pumping of air in and out of the lungs by the chest movements
50
usual among vertebrates. Moreover the lungs of turtles may be

forcibly emptied or filled by activities unrelated to respiration per
se; the lungs are the only compressible or expansible organs with
in the shell, so that when the turtle withdraws its head and limbs,
air has to be expelled from the lungs to make room. A turtle thus
may be obliged to exist for prolonged periods with its lungs almost
empty if it is subjected to a disturbance which requires it to retract
its extremities for a protracted period. Fortunately it has evolved
a number of physiological mechanisms which enable it to tolerate
such situations.
When one holds one’s breath, the circumstance which eventu
ally forces one to start breathing again is not so much shortage of
oxygen as an excessive level of carbon dioxide in the lungs; most
vertebrates cannot tolerate a high C02 pulmonary or haemal con
centration, and the presence of such is a strong stimulus to rapid
respiration. Turtles, however, are enabled to tolerate high C02
levels, whose immediate result is normally an increase in the acid
ity (or decrease in the pH) of the blood, by means of a buffering
system of bicarbonate ions, haemoglobin, and serum proteins in
the blood which resist pH changes. In other words, the blood can
contain more COa per volume at a particular pH than can that of
other organisms.
Various other modifications assist the turtle in maintaining
life while subject to the respiratory constraints mentioned above.
For one thing, the turtle is capable of filling and emptying its lungs
more completely than most vertebrates. This means that when a
turtle is suddenly obliged to withdraw after a period of active res
piration, it will initially have very little carbon dioxide in its lungs.
It is also significant that turtle meat is often very red. This is
caused by the high myoglobin content of the muscle, which means
that the turtle can store significant amounts of oxygen not only in
the haemoglobin in the blood but also in the muscular myoglobin.
Moreover, it will be recalled that many organisms can incur a so-
called ‘oxygen debt’—activity can be sustained for brief periods at
a level that requires more oxygen than the lungs can supply. In the
ensuing rest period, this ‘oxygen debt’ is paid off by panting, or
breathing more rapidly and deeply than such a resting phase
would normally require. This important ability is made possible
by an energy-producing reaction in the muscles by which glycogen
is converted to lactic acid without the requirement of oxygen.
However, lactic acid is somewhat toxic and must be burned off be
fore long by rest and hyperventilation. Turtles can tolerate much
higher levels of lactic acid than most organisms, and Dr. Daniel
Belkin has even demonstrated that at least one turtle species
(Stemotherus minor) may be able to depend indefinitely on this
anaerobic glycolysis and thus not require free oxygen. Moreover,
51
www.Ebook777.com
the heart muscle of many turtle species can operate anaerobically
for prolonged periods, and this phenomenon must be an important
factor in the ability of turtles to hibernate under water.
It should be mentioned, of course, that the ability to go for long
periods without respiring is not only useful to a terrestrial turtle
which retracts its extremities when disturbed, but it also enables
aquatic turtles to dive and remain submerged for prolonged peri
ods. The submergence time is lengthened still further by the slow
ing of the heart, or tachycardia, which occurs when aquatic turtles
dive, and which immediately lowers the body’s demands for oxy
gen. Moreover, many turtles can extract dissolved oxygen from
the water in which they are submerged. This phenomenon is un
derstandably more effective at low temperatures, when the crea
ture’s oxygen requirements are lowered and the solubility of oxy
gen both in the aquatic medium and in the animal’s body fluids is
increased. Various surfaces are used for oxygen transport in aqua
tic turtles, among which the most important are the thin papillose
skin of the kinostemids and chelydrids and the buccal lining of sea
turtles, emydids, trionychids, and dermatemyids.
Nevertheless, despite all of these mechanisms which minimize
the need for deep, regular breathing in turtles, turtles must still
have a mechanism for moving air into and out of the lungs, and it is
not immediately obvious how this is done, considering the confines
of the shell. It has been shown experimentally that the pressure in
side the lungs changes somewhat with every heartbeat and every
movement of the limb girdles, limbs, and neck, and these effects
may well allow sufficient air to move into and out of the lungs to
sustain certain types of moderate activity. However, I was impres
sed once after climbing a 3500 foot volcano in the Galapagos Is
lands to find that giant tortoises at rest on the rim of the crater
were breathing faster than I was. This active respiration is carried
out by means of a membranous sling across the rear part of the
shell, behind the lungs. This sling may be pulled forward, com
pressing the lungs and causing exhalation, by means of the trans
verse abdominal muscles, which enclose the posterior end of the
visceral cavity. The sling may be pulled back, expanding the lungs
and causing inhalation, by means of other muscles, the oblique
abdominals. Movements of these muscles also result in a pumping
in and out of the soft skin around the base of the hind limbs. There
are muscles at the anterior end of the lungs which also have respi
ratory functions—the diaphragmatic, used in exhalation, and the
testocoracoidials, used in inhalation—and which also, through
their attachment to the coracoids, produce a swinging movement
of the pectoral girdles when they contract. When a turtle is on land,
much of the work of inhalation may be carried out passively by the
weight of the relaxed, drooping viscera and limb muscles, which
52
produce a slight negative pressure in the lungs. Conversely, a tur

tle whose shell is submerged in water, with only the nostrils above
the water, can exhale passively under hydrostatic pressure and
only has to inhale actively. These mechanisms lessen the muscu
lar exertion of breathing, and in turn reduce the oxygen require
ments of the animal. When the necessity arises, of course, both in
halation and exhalation may become active, in water or on land.
The respiratory movements of sea turtles have not been prop
erly studied. However, the extremities of these turtles are non-re-
tractile, so incidental filling or emptying of the lungs as occurs
when other turtles retract or extend their extremities does not take
place. On the other hand, the chest is slightly expansible, since the
bridges are flexible, and some flexion is also possible along the
mid-line of the plastron. When a sea turtle is turned on its back on
land, it appears to hold its breath with lungs full for much of the
time, but to exhale and immediately re-inhale, with caving-in and
re-expansion of the chest at frequent intervals. This would con
trast with the normal respiratory rhythm in snakes, in which the
lungs at rest are about half filled, and during active respiration are
emptied, completely filled, and finally half emptied again to re
turn them to the original condition. Many turtles when at rest have
been observed to breathe normally for part of the time and then to
suspend breathing completely, sometimes for many minutes. Tur
tles can even survive for periods of hours in an atmosphere of pure
nitrogen. It would appear that under normal conditions the turtle’s
body contains sufficient oxygen to carry it through a protracted
period of complete oxygen deprivation.
The throat pulsations seen in many turtle species are appar
ently not concerned with respiration, but with pumping air over
the olfactory surfaces in order to ‘sample’ the environment.
DIGESTIVE AND EXCRETORY SYSTEM
The digestive system essentially follows the typical vertebrate
pattern. The esophagus, which in the marine species has a densely
spinose lining, joins the stomach at a point just dorsal to the left
atrium of the heart. The stomach is thick and curved and is attach
ed throughout its concave side to the left lobe of the liver by means
of the gastrohepatic ligament. The liver itself is very large in tur
tles. The duodenum begins just dorsal to the bridge of the liver and
is attached to the right lobe of the liver by means of the hepatoduo
denal ligament. Both of these ligaments are mesenteric in form.
The pancreas lies along the duodenum and discharges into it by
means of the pancreatic duct. The duodenum also receives bile
through the bile duct from the gall bladder, which is located on the
dorsal side of the right border of the liver. The small intestine,
which forms several curves, is attached to the midline of the body
53
www.Ebook777.com
by means of the dorsal mesentery. The large intestine, or colon, is
stout and relatively short; it is connected to the dorsal midline of
the shell by means of another mesentery, the mesocolon. The
spleen is located in the mesocolon. The colon terminates postero-
ventrally in the cloaca and anus.
The urinary bladder is typically thin-walled and bilobed. It re
ceives urine via the ureters from the kidneys and discharges pos
teriorly into the cloaca. The chemical and physical nature of the
nitrogenous waste product depend upon the abundance of water in
the habitat. It is known that some highly aquatic species (e.g. Che-
Ionia mydas) excrete considerable amounts of ammonia ( NH3), a
highly soluble and toxic material that requires the passage of
) arge volumes of water to prevent the formation of toxic concen-
1 rations in the body. Many semi-aquatic species, like many mam
mals, excrete urea ( NH2CONH2 ), a soluble and relatively non
toxic compound requiring some water for its elimination. How-
«ver, in those tortoises from xeric habitats for which water conser
vation is at a premium, urinary water loss cannot be tolerated and
the main nitrogenous excretory product is virtually insoluble uric
acid ( C5H4O3N4 ), which is non-toxic and is passed in the form of a
mass of white crystals. Such tortoises also use the urinary bladder
as a storage chamber for resorbed water; large tortoises can con
tain remarkable volumes of almost limpid water, and this drink
ing-water supply saved many of the early Galapagos explorers
from the ravages of thirst.
CIRCULATORY SYSTEM
The chelonian heart is normally considered to be three-cham
bered, in that the atria are paired but the single muscular ventri
cle is incompletely divided. The ‘beat’ of the heart originates in a
fourth chamber, the sinus venosus, which forces the deoxygenated
blood from the venous system into the right atrium through an
opening guarded by the sino-atrial valve. Contraction of the right
atrium squeezes blood through the atro-ventricular opening, again
guarded by a valve, into the right side of the ventricle, where a
partial spongy partition prevents complete admixture with the
blood in the left side of the ventricle. Blood from the right side of
the ventricle passes into the pulmonary artery, which divides and
conducts blood to the two lungs where it is oxygenated. The oxy
genated blood is returned to the left atrium of the heart via the pul
monary veins.
Blood from the left atrium is conducted to the left side of the
ventricle, whence it passes into the conus arteriosus and into the
great arteries. Three main arteries leave the conus in addition to
the pulmonary—the brachiocephalic and the two aortic arches.
54
The brachiocephalic artery divides into the two subclavian ar

teries, and each of these soon gives off a common carotid which
passes along the neck and finally forks into the internal and exter
nal carotids, conveying blood to the brain and facial areas respec
tively. The brachiocephalic also gives rise to the small coronary
arteries which supply the muscle of the heart itself with blood,
while small arteries branching from the subclavians supply blood
to the thyroid gland.
Each subclavian sends out several arteries to the underside of
the neck, the largest of which (the ventral cervical) has branches
to the esophagus and the thymus. The dorsal cervicals also branch
off the subclavians on each side. The anterior shoulder region re
ceives blood from the anterior suprascapular arteries, also bran
ches of the subclavians. Finally, an axillary artery branches off
the subclavian on each side, between the ventral cervical and the
anterior suprascapular, and this in turn gives rise to the vertebral
artery, the brachial artery, and the marginocostal artery. The
vertebral in its turn sends intercostal arteries between the dorsal
ends of the costal bones, which anastomose laterally with the mar
ginocostal artery. The brachial artery supplies blood to the front
limb.
The two aortic arches curve round behind and above the heart
and gullet, where they re-join to form the dorsal aorta. The dorsal
aorta continues, as the caudal artery, to the tip of the tail and
sends out many important branches to the viscera and hind limbs.
These include (i) the gastric artery (to the cardiac region of the
stomach, where it bifurcates into ventral and dorsal parts to sup
ply the inside and outside curvatures of the stomach respective
ly); ii) the coeliac artery, which branches into the anterior and
posterior pancreaticoduodenal arteries, which between them sup
ply the pyloric end of the stomach, the liver, pancreas, duodenum,
and, via the cystic artery, the gall bladder; iii) the anterior mesen
teric artery, which forms numerous branches radiating through
the mesentery to the small intestine; iv) the posterior mesenteric
artery, which conducts blood to the large intestines and cloaca; v)
the genital (ovarian or spermatic) artery; vi) the renal arteries
(two or three to each kidney); vii) the epigastric artery, of which
branches convey blood to the carapace, the marginocostal artery,
and the pelvic muscles; viii) the common iliac artery, which div
ides into internal and external iliacs. The internal iliac, which
gives rise to the haemorrhoidal artery, supplies the reproductive
organs, pelvic region, bladder, and large intestine. The external
iliac branches into the femoral artery (to the pelvic area and
thigh) and the sciatic artery (to the hind leg).
It should be noted in passing that the arterial branches even
within a single turtle species are highly variable, while the varia-
55
www.Ebook777.com
www.Ebook777.com
Venous system of an emydid turtle, Emys orbicularis. Redrawn from Bojanus, 1819.
tion that exists within the order as a whole has yet to be evaluated.
When conducting a comparative anatomy laboratory at the Uni
versity of Florida some years ago, I was hard put to explain why
even the great vessels of the heart of the Pseudemys scripta tur
tles being dissected by the class were so much more variable than
such trivial features as the stripes on the side of the neck.
Blood from all parts of the body except the lungs is returned to
the heart via the sinus venosus, which receives blood from the an
terior parts of the body through the precaval veins, and from the
posterior parts through the postcaval and the left hepatic vein. The
precavals receive blood from the head via the internal and
external jugular veins and their branches (the esophageal vein
drains into the internal jugular, and the brachial and axillary
veins, from the forelimb, drain into the external jugular). The pre
cavals also receive blood from the vertebral, scapular, subscapu
lar, marginocostal, and thyroid veins.
The most prominent veins in the abdominal region are the
paired abdominal veins, which arise on either side of the pelvic
girdle and disappear into the liver. Blood from the various muscu
lar and skeletal structures of the rear part of the body drains into
the abdominals via numerous veins—the caudal and cloacal veins
from the tail region, the crural and femoral veins from the hind
legs, the marginocostal veins from the sides of the shell, the pec
toral veins from the chest, the pericardial veins from the heart, the
pelvic vein from the pelvic area, the lipoidal vein from the abdomi
nal fat pad, and the vesicle vein from the bladder. Pressure in the
two abdominal veins is equalized by means of the short transverse
abdominal.
At their posterior ends the abdominal veins are connected via
the external iliacs with the renal portal veins, which thus offer an
alternative passage back to the heart through the kidneys. The
renal portals receive additional blood from the epigastric vein,
which drains the side of the shell; the hypogastric veins from the
rectum, cloaca, and the male genitalia; and the vertebral veins,
which conduct blood from the intercostal veins. The blood entering
the kidneys is filtered free of soluble waste products, then the
cleansed blood passes to the heart through the renals and the post
caval.
Venous blood from the digestive areas—the stomach, small in
testine, pancreas, spleen and gall bladder—conveys its absorbed
nutrients to the liver by means of the hepatic portal vein. The
hepatic portal passes along the rear of the liver from left to right,
receiving blood from the abdominals as well as from the gastric,
anterior and posterior pancreatic, cystic, duodenal, splenic, and
inferior and common mesenteric veins, terminating in the liver
sinusoids. Blood is drained from the liver by the hepatic veins into
58
the postcaval and back to the sinus venosus; the left hepatic enters
the sinus venosus directly.
NERVOUS SYSTEM
The turtle nervous system is not strikingly deviant from the
standard vertebrate pattern. The brain is covered by the membra
nous pia mater, outside of which is the thicker dura mater; the
brain is well protected by the skull roof, the large temporal mus
cles, and the thick braincase. The brain itself has a number of con
spicuous paired swellings; from the front, these are the olfactory
sacs, the olfactory bulbs, the cerebral hemispheres, and the optic
lobes. Median structures visible in dorsal aspect include the pineal
body, or epiphysis, located between the cerebral hemispheres and
the optic lobes; the cerebellum between the optic lobes; and the
choroid plexus between the cerebellum and the medulla oblongata.
The medulla oblongata is the posteriormost part of the brain and
tapers into the spinal cord. The function of the pineal body in
higher vertebrates is still somewhat obscure, but in certain primi
tive reptiles (e.g. the tuatara and the monitor lizards) it is light-
sensitive and is located just below an open perforation in the roof
of the skull. No turtle has a perforation in this area, but it is per
haps significant that the leatherback turtle (Dermochelys) has an
extremely thin spot in the skull roof above the pineal, and that the
skin over this thin spot is unpigmented. Perhaps there is some
residual light-sensitivity in the pineal of this species. The cerebel
lum has the function of coordinating the neuromuscular mecha
nism of the body. The choroid plexus is simply an area of the brain
where the roof of the brain is fused with the pia mater, and the re
sulting tela choroidea is thrown into a pattern of extensive folds.
Structures only visible in ventral aspect include the optic
chiasma beneath the cerebral hemispheres (whose function is to
convey optic impulses from each eye to the opposite side of the
brain, and in those animals with binocular vision to permit decus
sation or crossing-over of nerve fibers to take place to facilitate
visual integration), and the infundibulum and pituitary gland (hy
pophysis) just behind the optic chiasma. The pituitary is import
ant as the key endocrine gland of the body.
The brain and spinal cord are hollow, fluid-filled structures.
The various enlarged areas of the brain contain corresponding en
larged hollows known as ventricles; Ventricles I and II are located
in the two olfactory bulbs; each connects with the median Ventri
cle III via a Foramen of Monro. Ventricle III connects with Ventri
cle IV (located in the cerebellum) via the Aqueduct of Silvius.
Turtles have twelve pairs of cranial nerves. The anteriormost
(the olfactory nerves) connect the olfactory sacs to the olfactory
bulbs, and are associated with the sense of smell. The optic nerves
59
www.Ebook777.com
convey visual images from each eye to the ventrally-rooted optic
chiasma (although in general purely sensory nerves have a dorsal
root, purely motor nerves a ventral root, and nerves with both
these functions both dorsal and ventral roots). The oculomotor
nerve branches from the floor of the midbrain and innervates the
superior, inferior, and anterior rectus muscles and the inferior
oblique muscle, all of which function in movements of the eyeball.
The trochlear nerves (IV) branch from the dorsal side of the mid
brain and innervate the superior oblique muscles. The name
‘trochlear’ derives from the pulley-like arrangement over which
the tendon involved with this movement passes in man.
The trigeminal, or fifth, nerve is a thick and complex structure
which derives its name from its three branches. It originates from
the middle of the ventral wall of the medulla and connects with a
ganglion, or swelling, from which the three branches (the ophthal
mic, maxillary, and mandibular branches) pass one above
another. This nerve has both motor and sensory functions; the
branches named innervate the tissues around the eyes, the face,
and the lower jaw respectively.
Nerve VI, the abducens, branches from the midventral line of
the medulla and innervates the posterior rectus muscle of the eye.
The name abducens derives from its action of drawing the eyeball
away from the forward direction.
Nerves VII and VIII, the facial and auditory nerves, arise
together from a point lateral to the root of the abducens. The facial
passes anteriorly through the inner ear capsule and innervates the
muscles which raise and lower the hyoid bone. This is the only
motor function of the facial nerve, since a turtle, having the skin
closely adpressed to the skull, has none of the minor facial muscles
present in mammals. The auditory nerve, purely sensory in func
tion, has branches to the cochlea and the semicircular canals and
vestibule (the acoustic and vestibular branches, responsible for
the functions of hearing and balance respectively). Nerve IX, the
glossopharyngeal, arises just posterior to the common root of the
facial and auditory nerves, and has both sensory and motor func
tions in the tongue, pharynx, and anterior hyoid area. Nerve X, the
vagus, is a large and important nerve with both motor and sensory
functions. It has several rootlets along the side of the medulla, and
reaches far back into the viscera (the name vagus literally means
‘wanderer’), while a branch of the vagus gives rise to the sympa
thetic or involuntary nervous system.
Nerve XI, the spinal accessory, arises in close conjunction
with the root of the vagus and has the function of supplying motor
impulses to the muscles of the neck. Finally, the hypoglossal nerve
arises near the mid-ventral sulcus of the posterior medulla and
extends forward to innervate the muscles of the anterior hyoid
horn and the tongue.
60
A pair of spinal nerves develops from each vertebra of the

neck and trunk. These have both dorsal (sensory) and ventral
(motor) roots. They are somewhat gathered together in the re
gions of the limb girdles, forming the brachial and lumbo-sacral
plexi respectively. The brachial plexus derives from the sixth, sev
enth, and eighth cervicals, plus a small branch from the first thor
acic, and gives rise to the dorsal radial, the ulnar, and the median
nerve to the foreleg. The lumbro-sacral plexus is derived from the
last three thoracic nerves and the two sacral nerves, and gives rise
to the three sciatic nerves of the hind leg, as well as to small nerves
which innervate various muscles and skin along the outer part of
the thigh.
REPRODUCTION
In many turtle species, the sexes are of almost the same size.
However among giant tortoises and alligator snapping turtles
males are much larger than females, while for many of the
smaller species females are larger than males. Sexual
dimorphism reaches its extreme in certain Graptevxys species,
e.g. G. barbouri, in which the males only reach a carapace length
of about 4.5”, but females may be over one foot in length. Copula
tion is virtually always carried out in a position in which the male
is mounted on the carapace of the female, and males of many spe
cies (particularly the more strongly domed forms) have a con
cavity in the plastron which assists in the maintenance of the
mounted position. Males typically also have a longer tail than fe
males, with a more distally located vent. Males of certain sea tur
tle species have recurved hook-like claws on their front flippers
which allow them to grip the front edges of the female’s carapace
and thus stabilize their position. A third holding point is provided
by the prehensile, spur-tipped tail. On the other hand, American
box turtles have such bulbous shells that the male cannot possibly
reach the front edge of the female’s shell, and because of the wide
posterior plastral lobe intromission can only be completed with the
male leaning right over backwards. Specially recurved claws on
the hind feet allow the male to grip the hind edges of the female’s
carapace and thus maintain his position. Many male mud turtles
(Kinostemon) have roughened patches on the inner surface of the
hind legs, which appear to have a clasping function.
Some turtle species have rather elaborate courtship rituals.
Males of certain Pseudemys, Chrysemys, and Graptemys species
stroke the face of the chosen female with a rapid fluttering motion
of the forefeet, which are held with the palms outward as the tur
tles face each other. Males of these species have three exceptional
ly long claws on the front feet which appear to be specifically adap
ted for this type of courtship. On the other hand, males of many
www.Ebook777.com 61
tortoise species beat their females into submission (or more prob
ably into a state of hormonal readiness) by withdrawing their
heads and crashing the front part of their shells into the sides of
their prospective mates. Various courtship rituals displayed by
other species are described under the accounts for the individual
species.
Most turtle species are normally completely mute, but some
produce surprising sound effects when mating. Perhaps the most
dramatic sounds are the loud groans produced by courting and
mating giant tortoises, which may be heard from hundreds of
yards away. Many other tortoise species vocalize when mating;
further details are given later.
In both sexes the gonads are paired. The female ovaries are
capable of great distension, and in a gravid turtle ready to lay they
may occupy a very substantial part of the total visceral mass. Cor
pora lutea are formed in the ovaries after ova have been dischar
ged, and it is sometimes possible to estimate the size of the last
clutch by counting the corpora lutea. A dissected female turtle
often shows large numbers of yolks of all sizes from a millimeter
or so up to the full size of the yolk of the shelled egg. Apparently the
albumin layer and the shell are laid down fairly quickly, but the
shelled eggs are sometimes retained for some time before laying.
Under unnatural captive conditions eggs may be retained for so
long that they either erode through the walls of the oviduct or be
come hyper-calcified, a condition that prevents development of the
egg by eliminating the normal porosity of the shell, and thus pre
venting the ingress of oxygen. In the female turtle, the oviducts,
urethra, and intestine all discharge into the cloaca. A clitoris may
be present in the ventral wall of the cloaca; this is similar in struc
ture to the male penis.
The turtle penis is completely contained within the cloaca
when in the relaxed state; its presence causes a distinct thickening
and prolongation of the pre-cloacal part of the tail. Under sexual
excitement, the penis is extruded through the vent, and with full
erection curves downward and slightly forward. The erection is
caused by tumescence in two spongy bodies, the corpus spongio
sum and the corpus fibrosum, in contrast to penile erection in
snakes and lizards which is essentially a process of evertion. The
chelonian penis is tremendously variable in shape, and analysis of
its structure has been suggested as a means of establishing natural
relationships within the cryptodiran families. However, the varia
tion even within a genus (e.g. Stemotherus) may be greater than
that between families (e.g. Cheloniidae and Dermochelyidae), so
that this character should be regarded as only a supplementary,
rather than a critical, one.
62
Penis of Chelydra serpentina rossignoni.
www.Ebook777.com
SEX RATIOS
The subject of sex ratios and sex determination in turtle
populations is a complex one meriting far more study than it has
received.Various reports on unequal sex ratios in turtle popula
tions have been made from time to time. S.F. Hildebrand, for
example, found that a group of 1300 diamondback terrapins
(Malaclemys) hatched and raised in captivity yielded only 19%
males. Yet F.R. Cagle in 1952 found that a natural population of 70
diamondbacks included 81% males! Whitfield Gibbons, in a 1970
paper in Researches on Population Ecology, examined published
reports on unequal sex ratios and concluded that all were artifacts
resulting either from a biased sampling technique or from
inclusion of immature females, exclusion of the smaller mature
females, or failure to recognize adult males.
Nevertheless, I am convinced that sex ratios of some natural
turtle populations differ substantially from equality. For example,
the relatively undisturbed green turtle population in the
Galapagos Islands apparently includes far more females than
males, according to all local people who have any contact with
turtles. This would also be a logical stratagem for a population of a
species subject to intensive predation at the egg and juvenile
stages; more females means that a given total population can
produce more eggs, and of course one male can fertilize several,
perhaps many, females. Moreover, some populations of the genus
Rhinoclemmys, particularly those in South America, appear to
have substantially more females than males. A series of 17 in
dividuals of R. melanostema examined by Federico Medem in
cluded only four males, while two series of R. diademata that I ex
amined in Colombia and Venezuela showed an even more deviant
ratio: ten specimens in the Maracaibo Zoo, presumably randomly
caught locally, included only one male, while ten mature
specimens in the collection of the Instituto La Salle in Bogota in
cluded only two males. Although I have not kept figures on this, the
numerous R. punctularia that I have examined in Guyana and
Surinam over the years also were almost all females.
Assuming, then, that certain turtle species do naturally show a
significant divergence from a sex ratio of equality that is neither
the result of selective sampling or differential mortality, it is
interesting to speculate upon its possible genetic origins. It has
recently been shown that one turtle genus (Staurotypus) shows
male heterogamety of the mammalian type—that is, an unequal
pair of chromosomes in males and all pairs equal in females.
However, the other three genera of the Kinostemidae—Kino-
sternon, Sternotherus, and Claudius —do not show male
heterogamety, and it can be assumed that the condition in
64
Staurotypus is a relatively recently and independently derived one

rather than a fundamental one. In fact, in most turtles the sexes
cannot be distinguished by gross chromosome morphology.
Recent data from several independent workers have yielded
fascinating insight—and raised many new questions—about turtle
sex determination. Studies at the Cayman Turtle Farm, a
commercial green turtle ranching facility on Grand Cayman
Island, have revealed that the sex ratio of series of green turtles
from particular areas and hatched in particular years may
deviate substantially from equality; batches of green turtles from
Tortuguero, Costa Rica, for example, in some years yielded 80% or
more females and in other years 80% or more males. Parallel
observations by C.L. Yntema of the State University of New York
on the snapping turtle, Chelydra serpentina, showed that
environmental condition of incubation, especially temperature,
had a profound effect upon the sexes of the hatchlings; eggs
incubated at either 20° or 30°C developed into females in 100% of
the cases; at 26°, 99% of the individuals were males; at 24°, 100%
were males. More males than females developed at incubation
temperatures of 22° and 28°C. Additionally, Claude Pieau in
France hatched Emys orbicularis and Testudo graeca under
different experimental conditions; in one experiment, 42 Emys
eggs were reburied in a situation fully exposed to the sun, at a
depth of between 5 and 12 cm; of 28 embryos that developed, 27
were male and one intersexual. Pieau found that in Emys eggs
incubated below 28° to 29° produced only male hatchlings, and only
females when incubated at temperatures above that range;
similarly, a ‘‘critical temperature” of between 30 and 31° was
found for Testudo graeca, below which all eggs yielded males and
above which all females. The ultimate sex ratio of a turtle
population, then, would appear to be a result of rather fortuitous
circumstances, namely the prevailing average incubation
temperature during the various seasons in which the standing
population hatched. Whether this seemingly chancy mechanism
has some subtle survival advantage over purely genetic means of
sex determination remains to be demonstrated; possibly turtle
populations contrive to exist in spite of it rather than because of it.
TURTLE EGGS
All turtle species reproduce by means of eggs, which are de
posited in (or occasionally on) the ground and abandoned once the
nest site has been suitably disguised. The eggs vary greatly in
number, size, shape, and texture. The variation in these para
meters is complex and interconnected, and each represents a bal
ance between several opposing environmental constraints and
65
www.Ebook777.com
selective factors. For example, the total number of eggs and the
size of each tend to be inversely related for turtles within a certain
size bracket, simply because the space within the shell available
for eggs is limited. In certain situations it may be desirable to pro
duce as many offspring as possible to increase the statistical
chances of survival in a highly exacting habitat. The sea turtles,
for example, may lay seven or more times in a season and fre
quently produce clutches of well over a hundred eggs. The two lar
gest sea turtle clutches on record, both laid by green turtles, each
contained 226 eggs. One clutch was laid in the Philippine Turtle
Islands in 1951, the other in Surinam in 1967*
At the other extreme are such species as the African pancake
tortoise (Malacochersus) and certain of the South American Rhi
no clemy s, which typically lay only a single large egg at a time.
Presumably this low reproductive potential is sufficient to main
tain population levels in the particular ecosystems in which the
animals live. Yet it is still obscure exactly why an animal should
evolve so as to realize less than its full reproductive potential,
since any individuals which have a tendency in that direction
would necessarily be the ones that have a diminished effect in pro
viding the genes for the next generation, other things being equal.
However, several possible partial explanations do present them
selves. For example, many turtle populations are relatively non
dynamic—that is, it is more important for an adult of such a popu
lation to survive for a long time and reproduce itself rather mod
estly than for it to exhaust itself producing large numbers of eggs
(none of which may ultimately produce breeding adults), and then
expire from this effort before the next breeding season—if this is
indeed the choice that presents itself.
It is possible that this is the mechanism that obtains among
Galapagos tortoises, for example. These tortoises normally lay
fewer than fifteen eggs, typically twice per year, while on some of
the more arid islands (such as Duncan) a nest may contain only
two or three eggs; yet sea turtles of comparable body size will lay
many hundreds of eggs in the course of a single season. Primordl-
ally, Galapagos tortoise populations were so abundant that the ul
timate limiting factor was the availability of food; perhaps a tor
toise seriously reduced its chances of surviving through the dry
season if it converted too great a fraction of its food intake into
eggs. It is possible that calcium availability may also be the limit
ing factor for egg production in some cases; female Duncan tortoi
ses have highly fenestrated bony shells, presumably because the
necessity of providing calcium for the egg shells extracts calcium
phosphate from the shell.
*Recently a hawksbill turtle nesting on Cousin Island in the
Seychelles laid 242 eggs.
66
Turtle eggs do not vary in size nearly as much as do the adult

turtles; the smallest turtle eggs are a little less than an inch in
maximum diameter, while the largest still measure less than
three inches in maximum diameter. This small size variation may
reflect the fact that extremely large or small adult size is a spe
cialization that would not be expected to appear in hatchlings and
consequently would not be fully reflected in the size of the eggs,
coupled with the fact that extremely small turtles tend to produce
relatively large offspring, because a truly diminutive hatchling
would have very limited survival prospects.
Many of the turtles which produce hard-shelled eggs in fact lay
the largest eggs which can fit through the hind opening of the shell,
between the pygals and the xiphiplastra. This limiting factor has
had interesting results in the case of the South American tortoise
Geochelone denticulata. This species has a heavy, thick shell, with
relatively narrow openings at front and back, presumably to dis
courage predation by jaguars and other large carnivores. The hind
margin of the plastron has a semicircular notch, and this notch is
just large enough to allow the passage of an egg, making allow
ance for the extra diameter of the distended tissues of the tail and
cloaca. However, this species reaches sexual maturity at about
twelve inches, while the largest mature females may be up to
thirty inches in carapace length. Large specimens lay eggs that
may have up to three times the volume of those laid by small,
younger females. Consequently the hatchling is larger when the
mother is larger, and presumably reaches a larger ultimate size
than the offspring of small mothers. So the unusually wide size
range of mature denticulata would tend to increase. It seems
likely that larger hatchlings have a better chance of survival than
small ones, so the tortoise has evolved the ability to lay the largest
eggs that are commensurate with the necessity of also maintain
ing a completely armored body. However, there is no shortage of
food or moisture in the rain forest inhabited by this species, and
consequently a female tortoise loses nothing by producing some
eggs, necessarily of smaller size, when it is still well short of its ul
timate body size. Moreover it may well gain something, since even
the small hatchlings from these early eggs will have some chance
of surviving to maturity.
Sometimes closely related species lay very different numbers
of eggs. For example, Chelonia depressa only lays about fifty eggs,
on average, while Chelonia mydas usually lays well over a hun
dred. An even more extreme case is that of Podocnemis unifilis
and P. expansa; according to Federico Medem, the former usually
lays between 15 and 25 eggs, while the latter may lay more than one
hundred. The most probable explanation is that in these cases the
less prolific species combines smaller body size with larger egg
67
www.Ebook777.com
size, so the total number of eggs that the animal can contain is
much smaller. However, in the case of the American gopher tor
toises of the genus Gopherus it is likely that a different mechanism
prevails. The Florida gopher, Gopherus polyphemus, usually lays
five or six eggs, though rarely as few as two or as many as seven.
The California desert tortoise, Gopherus agassizi, lays even
more—sometimes as many as thirteen. The Texas tortoise, G. her-
landieri, on the other hand, most commonly lays only one egg, less
often two, and only occasionally three or four. We have reasoned
earlier that infertility, or low fertility, is by definition an unlikely
characteristic for evolutionary selection. However, the ecologist
G. Maynard Smith has advanced a model, based on the controver
sial concept of group selection, which may be adapted to explain
the low fertility of the Texas tortoise.
Much of the habitat of the Texas tortoise consists of hillocks, or
lomas, rising out of a flat plain which, by virtue of its lack of shade
and food, is not entered voluntarily by tortoises, and only occasion
ally traversed involuntarily in times of flood. The tortoise popula
tions on the lomas may reach very high densities, and in times of
drought food may be the factor which limits population size. If, by
chance, the population on a particular loma were founded by tor
toises of only modest fertility, the population would be unlikely to
outstrip the available food supplies. However, a nucleus of highly
fertile tortoises would be liable to overpopulate the habitat; the
population would survive until all food was consumed, and then
would collapse. Consequently, if inter-loma migration were suffi
ciently rare, group selection would actually favor low fertility in
this case. On the other hand, we may be putting the cart before the
horse; perhaps the high population densities and relatively low
productivity of the habitat simply do not afford the female tortois
es sufficient nourishment to produce a lot of eggs, as we have pos
tulated may have been the primordial clutch-limiting mechanism
for Galapagos tortoises.
Turtle eggs vary in shape from spherical to markedly elon
gate. Basically, we would expect eggs to have a spherical shape
unless they have a particular reason not to be (birds’ eggs are
pointed at one end to lessen the likelihood of their rolling out of the
nest), and we find that all the biggest turtles—the sea turtles, the
giant tortoises, the snappers, soft-shells, matamatas, Podocnemis,
etc.—do indeed produce spherical eggs. An egg of this shape also
has the minimum possible ratio of surface to volume, and conse
quently will be favored when the eggs are deposited in an arid en
vironment and evaporative water loss becomes a problem. Thus
not only giant tortoises, but also medium-size ones and some of the
smaller ones lay eggs that are almost perfect spheres. On the
other hand, very small turtles or those which for various reasons
68
Hatchling of the common box turtle, Terrapene Carolina. Above, ventral view
showing the umbilicus; below, emergence from egg. Photos by R.J. Church.
69
www.Ebook777.com
have a very narrow separation between the hind margins of the
carapace and plastron are forced either to lay elongate eggs or
very small spheres; and selection usually favors the former, since
a tiny sphere simply could not produce a tortoise of adequate size
for survival. Among tortoises, for example, the diminutive forms
such as the various species of Homopus, Malacochersus, and
Psammobates lay elongate eggs, and so does Gopherus berlandieri
which has so narrow a posterior shell opening that the tips of the
anal scutes may almost touch the hind margin of the carapace.
Freshwater turtles of the family Emydidae also normally lay
elongate eggs, as do the Kinostemidae and Dermatemyidae.
Among the sidenecks (Pleurodira) both elongate and spherical
eggs may be found.
Turtle eggs shells may be either flexible and leathery or brittle
and well-calcified. Flexible eggs are laid by the sea turtles and
many emydids; on the other hand most other turtles lay hard-
shelled eggs. Flexible shells have the advantage of requiring less
calcium for their formation, and there is also no danger of their
cracking when they are dropped on to each other during oviposi-
tion. But eggs are laid on the surface (as are those of Geochelone
carbonaria in parts of its range, some Rhinoclemys, and some
Kinostemon), or those which are buried in the ground in arid
places (most tortoises), would dehydrate fatally if they had soft,
permeable shells. Others, such as those of Dermatemys, are laid in
such moist situations that a hard shell is needed to keep moisture
out, rather than in. Flexible shells are good osmotic membranes,
so that sea turtle eggs, for example, become dehydrated when ex
posed to salt water, while under normal conditions of natural in
cubation, they absorb fresh water through the shell so that the dent
present when they are laid is eliminated as the eggs develop.
Hard-shelled eggs have the liability of being brittle and easily
broken. Giant tortoises, which make relatively deep nests and lay
large, hard-shelled eggs, produce a gush of viscous mucus which
surrounds each egg as it is laid, so that the impact is cushioned as
the egg falls on its fellows. This mechanism, however, is easily
disrupted, and in captive situations it has been observed that
Galapagos tortoise eggs may actually crack as they are dropped
into the nest.
BACKGROUND READING
Ashley, L.M. 1962. Laboratory Anatomy of the Turtle. Wm. C.
Brown Co., Iowa. Pp. 1-48.
70
Auffenberg, W. 1966. The carpus of land tortoises (Testudininae).

Bull. Florida State Mus., vol. 10, no. 5, pp. 159-191.
Belkin, D.A. 1962. Anaerobiosis in diving turtles. Physiologist,
Washington 5: 105.
------------- 1964. Variations in heart rate during voluntary diving
in the turtle Pseudemys concinna. Copeia 1964: 321-330.
------------- 1965. Critical oxygen tensions in turtles. Physiologist,
Washington 8: 109.
------------- 1968. Anaerobic brain function: effects of stagnant and
anoxic anoxia on persistence of breathing in reptiles. Science
162: 1017-1018.
Bojanus, L.H. 1970. Anatome Testudinis Europaeae. Reprint Edi
tion, Society for the Study of Amphibians and Reptiles, Ohio.
Pp. i-vi, 1-178, 31 pis.
Carr, A.F. 1952. Handbook of turtles. Comstock publ. Assoc.,
Ithaca, N.Y. 542 pp.
Coker, R.E. 1910. Diversity in scutes of Chelonia. J. Morph. 21,
1-75.
Gadow, H. 1899. Orthogenetic variation in the shells of Chelonia. In
Wiley’s Zool. Results. Pt. Ill, 207-262.
Gaffney, E.S. 1972. An illustrated glossary of turtle skull nomen
clature. Amer. Mus. Novitates, No. 2486:-l-33.
------------- 1975. A phylogeny and classification of the higher cate
gories of turtles. Bull. Amer. Mus. Nat. His. 155(5): 387-436.
George, J.C. and R.V. Shah. 1954. The occurrence of a striated
outer muscular sheath in the lungs of Lissemys punctata gran-
osa Schoepff. J. Anim. Morph. Physiol., 1.1.13.
-------------------------- 1955a. Respiratory mechanism in the Chelo
nia. J. Anim. Morph. Physiol. 1.2.30.
-------------------------- 1955b. The myology of the chelonian trunk
and tail. J. Anim. Morph. Physiol., 2.49.
-------------------------- 1959. The structural basis of the evolution of
the respiratory mechanism in Chelonia. J. Anim. Morph. Phy
siol., 5.1.
Lumsden, T. 1924. Chelonian respiration. J, Physiol., 58: 259-266.
Lynn, W.G. and M.C. Ullrich. 1950. Experimental production of
shell abnormalities in turtles. Copeia No. 4: 253-262.
Medem, F. 1960. Datos zoo-geographicos y ecologicos sobre los
Crocodylia y Testudinata de los Rios Amazonas, Putumayo y
Caqueta. Caldasia, 8 No. 38: 341-351.
Mosimann, J.E. 1958. An analysis of allometry in the chelonian
shell. Rev. Can. Biol. 17: 137-228.
Newman, H.H. 1906. The significance of scute and plate ‘abnor
malities’ in Chelonia. Biol. Bull. 10: 86-114.
Pritchard, P.C.H. 1966. Occurrence of mesoplastra in a cryptodi-
ran turtle, Lepidochelys olivacea. Nature, 210, No. 5036: p. 652.
71
www.Ebook777.com
__________. 1971. Numerical reduction of bony plastral elements in
the kinostemid turtle Claudius angustatus. Copeia No. 1, pp.
151-152.
Ruckes, H. 1929. The morphological relationships between the gir
dles, ribs and carapace. Ann. N.Y. Acad. Sci. 31: 81-120.
Schumacher, G.H. 1954. Beitrage zur Kiefermuskulatur der
Schildkroten. I. Mitteilung. Wiss. Zeitschr., Univ. Griefs-
wald, Jahrgang 3, Math.-Naturwiss., pp. 457-518.
__________. 1955a. Beitrage zur Kiefermuskulatur der Schild
kroten. II. Mitteilung, Ibid., Jahrgang 4, pp. 501-518.
__________. 1955b. Beitrage zur Kiefermuskulatur der Schild
kroten. III. Mitteilung, Ibid., Jahrgang 4, pp. 559-587.
__________. 1956. Morphologische Studie zum Gleitmechanlsmus
des M. adductor mandibularis extemus bei Schildkroten. Anat.
Anz., 103: 1-12.
Shah, R.V. 1961. The occurrence of a pericardial extension of the
lung muscle in some turtles. Brit. J. Herpetology, 2 (12): 215-
217.
Thompson, J.S. 1932. The anatomy of the tortoise. Sci. Proc. Roy.
Dublin Soc., 20: 359-461.
Volker, H. 1913. Uber das Stamm-, Gliedmassen-, und Hautskelett
von Dermochelys coriacea L. Zool. Jb. Abt. Anat. 33. 431-552.
Williams, E.E. 1950. Variation and selection in the cervical central
articulations of living turtles. Bull. Amer. Mus. Nat. Hist., 94:
505-562, 20 figs.
__________and S.B. McDowell Jr. 1952. The plastron of soft-shelled
turtles (Testudinata: Trionychidae): a new interpretation. J.
Morph. 65: 383-406.
Zangerl, R. 1969. The turtle shell. In Biology of the Reptilia, Vol. 1,
London and New York. Pp. 311-339.
Zug, G.R. 1966. The penial morphology and the relationships of
cryptodiran turtles. Occ. Pap. Mus. Zool. Univ. Michigan, No.
647: 1-24.
72
Chapter 3
TURTLE EVOLUTION AND FOSSIL HISTORY
The Order Testudines has a long and distinguished history, and

in former geologic eras, turtles reached a far greater diversity
than they do today. The first unquestionable turtles are of Triassic
age, but despite the numerous primitive features of these early
representatives, they shed little light on the evolution of the Order
Testudines from its presumed cotylosaurian ancestors. Until
recently, it was customary to cite a reptile known as Eunotosaurus
africanus, from the Permian of South Africa, as a sort of ‘missing
link’ between the cotylosaurs, or stem reptiles, and the true turtles.
This creature is known from several incomplete fossils in the
British Museum, which show it to have been very small (only four
or five inches long), and to have eight pairs of expanded ribs, as is
typical of turtles today. However, certain critical parts of the
specimens, notably the skull roof, the neck, and the feet, are miss
ing, and it is now thought that Eunotosaurus was a deviant
cotylosaur that was not in the main line of testudinate evolution.
The ribs, although eight in number, are leaf-shaped and quite
dissimilar to the sutured rib-pleural combination bones of modern
turtles, and the fossils available show no traces of dermal armor.
Eunotosaurus africanus, once considered an ancestor to the turtles.
73
www.Ebook777.com
Turtles are customarily divided into three suborders: the now
entirely extinct Amphichelydia, and the living Pleurodira and
Cryptodira. The Amphichelydia are defined by various primitive
features, including the usually complete skull roof; the pterygoids
excluding the quadrates from the basisphenoid; the presence of
one or two pairs of mesoplastra in many genera; the constant
strong contact of the pelvis with the plastron; the non-retractile
neck with high spines on the posterior cervical vertebrae; the con
stantly non-uni ted cervical postzygapophyses; and the frequently
amphicoelous, never doubled cervical centra. Various of these
primitive features are retained in modem turtles; for example,
pleurodires have the pelvis fused to the plastron, while one of the
two living families retains a pair of mesoplastra. The cervical cen
tra are single in pleurodires. The complete skull roof is retained in
certain modem turtles in which the neck is not fully retractile;
these include all the sea turtles, Platystemon, Pseudemydura, and
Peltocephalus tracaxa. In other living species, the skull roof is
emarginated from behind (or, in the Chelidae, from below) to
varying degrees, but true fontanelles are never present.
AMPHICHELYDIA
The Amphichelydia, in addition to the primitive and uncertain
family Archaeochelydiidae, contains three superfamilies, the Pro-
ganochelyidea (previously known as the Triassochelyoidea), the
Pleurostemoidea, and the Baenoidea. The Archaeochelydiidae are
represented by a single species, Archaeochelydium pougeti (Ber-
gounioux 1958). The available material, reportedly from the Per
mian of France, is too incomplete to make a proper definition of the
family; it may not be chelonian, or even vertebrate.
The Proganochelyidea have a complete skull roof, a parieto-
squamosal arch, two pairs of mesoplastra, free cervical ribs, the
last cervical neural arch fused to the nuchal bone, and amphicoe
lous cervical vertebrae. This combination definitely establishes
the Proganochelyidea as the most primitive of all turtles. Two
families (sometimes considered only as subfamilies) are recogniz
ed, the Proganochelyidae and the Proterochersidae; they are dif
ferentiated by the pelvis being fused to the plastron in the former
and not in the latter. The Proganochelyidae are known from two
species, Triassochelys dux and Proganochelys quenstedti. These
were both from the Triassic of Germany. In such early turtles the
scute patterns had not been stabilized and, although four or five
broad vertebrals and five pairs of costals are recognizable, the
marginals are very numerous and liberally supplemented with in
framarginals and supramarginals. The bones of the primitive
shoulder girdle, although partially merged with the plastron, are
74
75
www.Ebook777.com
the acromion is short. This last subfamily is sometimes given full
family status. Another family, the Aperotemporalidae, may belong
here, but being known only from skull fragments, a definite
disposition cannot yet be made.
The Pleurosteminae are known from the Jurassic and Creta
ceous of Europe and North America. They tended to have elongate,
flattened, strongly tuberculate shells and frequently cruciform
plastra. A number of genera are recognized, including Pleuroster-
non, Platychelys, Glyptops, Archaeochelys, Neomichelys, Platy
chelydites, Probaena, Protochelys, and Trachydermochelys.
The Desmemydinae is based on the single species Desmemys
bertelsmanni, a moderately large form from the Cretaceous of
central Europe. Desmemys was clearly adapted for marine life
and had the extensive intercostal and midplastral fontanelles
typical of modem sea turtles.
The single genus of the Kallokibotiinae, Kallokibotion from the
Upper Cretaceous of Europe, apparently was adapted for life in
coastal marshes and lagoons. The skull was completely roofed and
was very deep posteriorly. The margins of the shell are not
preserved in the available material, but precostal and In
framarginal scutes were present. Two or three species are known;
it is possible that the species magnificum is a synonym of bajazidii.
The Plesiochelyidae are represented by several genera from
the Jurassic and Cretaceous of Europe and Asia, including Plesio-
chelys, Craspedochelys, Tienfuchelys, Parachelys, Hylaeochelys,
and Scutemys, though it is possible that some of these genera will
be referred elsewhere when better material is available.
The Thalassemydidae were primitive sea turtles from the late
Jurassic and Cretaceous of Europe; it seems very likely that they
are descended from the Plesiochelyidae. They had open fon
tanelles between the rib-ends and at the sides of the plastron, and
as with modem sea turtles, the carapace and plastron were not fus
ed at the bridges. The neural bones were reduced in number and
size in many species. In the type genus Thalassemys, the en-
toplastron is apparently absent, the epiplastra are rudimentary or
lacking, and a large median plastral fontanelle Is present. On the
other hand, in the related genus Idiochelys the plastron was large
and had no median fontanelle. In many species of this family a
notch is present in the carapace behind the head. Other genera
than those mentioned include Eurystemum, Acichelys, Chelonid-
es, Hydropelta, Cirnochelys, Chelonemys, Pelobatochelys,
Pygmaeochelys, Sontiochelys, Tropidemys, and Jaxartemys.
The Aperotemporalidae were most probably marine forms
related to the Thalassemyidae. At the present time, they are only
known from skull material. The family is characterized by the
elongate skull, with shortened facial region and reduced temporal
76
region. Because of the superficial similarity to the skull of the liv

ing softshell Chitra, one of the genera is named Chitracephalus;
the other genus is Aperotemporalis. The family is known from the
Jurassic and Cretaceous of Europe and Egypt.
The Baenoidea are traditionally assumed to include five
families, the Baenidae, Eubaenidae, Macrobaenidae, Neurankyli-
dae, and Meiolaniidae. However, a recent revision by Eugene
Gaffney grouped the Baenidae, Eubaenidae, and Neurankylidae
together as one family. According to Gaffney’s revision, the family
Baenidae includes five subfamilies: the Trinitichelyinae, containing
only Trinitichelys hiatti; Hayemydinae, containing only Hayemys
latifrons; Eubaeninae, containing Plesiobaena antiqua, P. puto-
rius, Eubaena cephalica, and Stygiochelys estesi; Palatobaeni-
nae, containing only Palatobaena bairdi; and Baeninae,
containing Baena arenosa and Chistemon undatum. Compsemys,
Neurankylus, Boremys, and Thescelus, although included with the
Baenidae, are of uncertain subfamily, being known only from shell
material. Gaffney considered the Meiolaniidae and the Macrobae
nidae not to belong to the Baenoidea, but he did include the family
Glyptopsidae, containing the single species Glyptops plicatulus.
Geologically, the Glyptopsidae occur in the Upper Jurassic
and the Baenidae in the Cretaceous, Paleocene, and Eocene. The
Baenidae are characterized by the extensive skull roof, single pair
of mesoplastra, and short, broad head, the length and width of
which were usually almost equal. The earlier Baenidae had
amphicoelous cervical vertebrae, but formed central articulations
are found in the later members. Baena itself was once assumed to
include eighteen species, but these are all now synonymized with
the highly variable Baena arenosa. The bewildering variability of
this species is especially evident in the scute arrangements on the
anterior part of the carapace. The Glyptopsidae differ from the
Baenidae in having very elongate skulls, narrow triturating sur
faces with no medial expansions, the basisphenoid extending the
length of the pterygoids, the probably present epipterygoid, the
incisura columellae auris widely open posteroventrally, and the
last vertebral scute barely enclosed posteriorly by tapering mar
ginal scutes (either completely enclosed or widely open in the
Baenidae).
Chengyuchelys baenoides, from the Upper Jurassic and Lower
Cretaceous of China, has been considered to be a baenid, but
according to Gaffney the described material lacks distinctive bae
nid features. Chengyuchelys had a flattened neural region, no
plastral fontanelles, a rhomboidal entoplastron, and narrow meso
plastra.
The large species Macrobaena mongolica, from the Eocene of
Naran Bulak, Mongolia, had an oval carapace with narrow, rec-
77
www.Ebook777.com
tangular neural bones, recostal scutes, no mesoplastra, a large,
kite-shaped entoplastron, and very narrow epiplastra. The
plastron was small but had broad, fully-sutured bridges. The first
cervical is procoelous, the fifth amphicoelous. This is the sole
species of the family Macrobaenidae.
The Meiolaniidae were very large, specialized turtles related
to the Baenidae. Three genera are known: Niolamia, with the
single species N. argentina, from the Upper Cretaceous of Pata
gonia; Meiolania, with two species, M. oweni and M. platyceps,
from the Pleistocene of Eastern Australia, Lord Howe Island, and
Walpole Island; and Crossochelys, with the single species C. cor-
niger from the Eocene of Argentina. Meiolania oweni was an enor
mous turtle with a massive, knobbed skull whose laterally-placed
conical horns had a span of nearly two feet in the largest
specimens. The survival of such a bizarre species as late as the
Pleistocene is truly remarkable, especially when one remembers
that the very similar Niolamia, whose horns were more flattened
and which had a raised flange-like frill at the rear of the skull, was
a late Mesozoic form from the other side of the Pacific Ocean. The
tail of Meiolania was encased in bone, but it is not known why the
animal required such extensive armor. It appears that Meiolania
was a terrestrial form and, having been present on certain remote
oceanic islands until quite recently, may have been a sort of
ecological equivalent to the giant tortoises of today.
Skull of Meiolania, a
Pleistocene Australian
horned turtle.
PLEURODIRA
Of the two extant families of this suborder, the Chelidae,
represented today by several genera In the Australasian and South
American regions, is poorly represented in the fossil record, with
no fossils known from outside the area where the family is still
found. Among the few fossil genera known are Parahydraspis,
from the Pleistocene of South America, and Pelocomastes from
the Pleistocene of Australia. The distribution of the family thus
78
parallels that of the Meiolaniidae. On the other hand, the Pelome-

dusidae, which today are confined to South America, Africa, and
Madagascar, have a long and rich fossil history extending from the
Cretaceous throughout the Tertiary. The family is defined by the
laterally retractile neck, presence of a single pair of mesoplastra,
the absent nuchal scute, the present or absent vomer, and the
biconvex second cervical centrum. Certain fossil forms from the
Cretaceous to Miocene of North America are included in a
separate subfamily, the Bothremydlnae, distinguished by the
presence of the vomer and the deep pit In the triturating surfaces of
the maxillae and mandibles. A different division separates the liv
ing genus Pelusios from other pelomedusids because of the large,
mesially contiguous mesoplastra and anteriorly hinged plastron.
This latter classification, establishing the subfamilies Pelusiinae
and Pelomedusiinae, may be more logical, since the plastral dif
ference correlates with a much more emarginated skull roof in
Pelusios. On the other hand, the presence or absence of the vomer
is not as good a character as might be anticipated, since
Podocnemis vogli, which in all other respects is a typical
Podocnemis, lacks the vomer.
In North America, two fossil pelomedusid genera, Bothremys
and Taphrosphys, are considered valid, while two others, Ambly-
peza and Naiadochelys, are represented by such fragmentary
material that definite allocation cannot be made. Bothremys is-
known from quite abundant material from the Cretaceous of North
America referable to the species B. barberi (until recently known
as Podocnemis alabame). Two other species, B. cooki and B.
miocenica, have been described on the basis of fragmentary
material (a skull and mandible, and the anterior part of a plastron
respectively). The latter species, from the Miocene of New Jersey,
represents the only known Tertiary pelomedusid from North
America. Bothremys had a very Podocnemis-like shell, with six or
seven neural bones and widely separated mesoplastra, but the
skull differed from that of Podocnemis in being shorter and broad
er, in having more separated orbits and a more dorsally located
facial region, and especially in the presence of large paired pits in
the triturating surfaces, which may have functioned in holding
hard-shelled food organisms.
Taphrosphys is characterized by the presence of a large inter-
gular scute which completely separates both gulars and humerals
and partially separates the pectorals. In one of the eight or so
known species (T. molops), however, the intergular is dispatched
posteriorly so that the gulars meet in front of it, giving a configura
tion similar to that of the modern chelid Chelodina. With the excep
tion of T. olssoni from the Eocene of Peru, this genus is only known
from certain deposits, ranging from Cretaceous to Miocene age,
79
www.Ebook777.com
from New Jersey. It appears probable that these early pelomedu-
sids occupied a marine habitat.
Amblypeza entellus, from the Eocene of New Jersey, is closely
related to Taphrosphys, but had an extremely broad carapace
which included a nuchal scute—a character not observed in any
other pelomedusid. Naiadochelys is based on some highly frag
mentary and questionable material from New Mexico, and further
specimens from the Late Cretaceous of Patagonia reportedly re
ferable to two species, N. patagonica and N. major.
The subfamily Podocneminae itself, still represented by seven
South American and one Malagasy species now referred to three
genera, is one of the few Mesozoic reptilian groups which survives
today. Fossil Podocnemis have been reported from Egypt, Malta,
Germany, England, India, Congo, Venezuela, and Brazil, from
deposits of Cretaceous, Paleocene, Eocene, Oligocene, and
Miocene age.
Three fossil Podocnemis species have been described from
Brazil—P. harrisi, P. brasiliensis, and P. elegans. P. elegans,
from the late Cretaceous of the Bauru Formation, is based on a
good shell and a skull of a different individual, and is the oldest
known Podocnemis species. The other two species are both dubi
ous, since the available material, although clearly pleurodiran, is
only questionably referable to Podocnemis. Another species from
Peru, P. bassleri, is based on a well-preserved skull from Tertiary
deposits of unknown age. The skull is almost identical to that of the
living P. expansa.
A third South American species, P. venezuelensis, from the
mid-Pliocene of Venezuela, has recently been investigated in some
detail by Roger Wood. This species differs from other Podocnemis
in totally lacking neural bones. While this may be considered so
fundamental a character that It should warrant generic status, the
species is in other respects so typical of Podocnemis, and the
presence or absence of neural bones in the related family Chelidae
Is so variable even within a species, that the character is not as im
portant as might be thought.
It should be mentioned that a strange, almost perfectly cir
cular turtle, from the Eocene of Tunisia, with widely separated
mesoplastra and no neural bones has been declared the type
specimen of Eusarkia rotundiformis, which was placed by
Bergounioux in a new family, the Eusarkiidae. However, rotun
diformis could probably be better considered as a species of
Podocnemis.
A fossil pelomedusid is known from the Oligocene of Puerto
Rico, but the material is too fragmentary to be assigned generical-
iy-
Other fossil Podocnemis species that have been described in
clude P. aegyptiaca and P. bramlyi, from the Miocene of Egypt, P.
80
fajumensis from the Oligocene of Egypt, P. stromeri from the Eo

cene of Egypt, and P. bowerbanki from the Eocene of England.
These species are differentiated principally by the relationship be
tween the sulci and sutures in the anterior lobe of the plastron.
Podocnemis indica, from the Eocene of India, is in fact not defi
nitely referable to Podocnemis, or even to the Pelomedusidae,
though the neural pattern suggests that it might be a pelomedusid.
A species of Podocnemis from the Pliocene of northwestern Kenya
has recently been described.
In 1976, Roger Wood described a new, gigantic fossil pelo
medusid from the Huayquerian (probably Pliocene) of Falcon
State, Venezuela. Wood claimed that this species, which he named
Stupendemys geographicus, is the largest turtle that ever lived;
the two specimens collected had respective carapace lengths of 218
and 230 cm (7’2” and 7’61/£”), and these were straight, midline
lengths, the total shell length being about seven or eight inches
more since the nuchal area is strongly indented. This is far longer
than the living leatherback turtle, which is not reliably known to
exceed 6 feet (183 cm). Archelon ischyros, a protostegid turtle that
is usually considered to be the largest species of all time, was not
really as large as often supposed; the type specimen has a cara
pace length of only 193 cm (76”), though the shell was much wider
than that of Dermochelys and the enormous, elongate skull prob
ably gave the animal a total length of at least nine feet. The largest
known land tortoises, Geochelone atlas and G. crassiscutata, did
not exceed a shell length of six feet.
Stupendemys was not only enormous, but also had an extreme
ly thick shell. The generic characters include the strong nuchal
indentation (possibly reflecting a massive, armored head similar
to that of Platystemon or Macroclemys, but the skull of Stupen
demys is still unknown) and the presence of a curious thickened,
upturned area on the front of the nuchal bone. The shell was rather
low, and the humeri and femurs extremely squat and massive. The
habits of this enormous turtle are of course a matter of pure con
jecture, but it must surely have lived either in rivers of great size
or in the sea. It is unfortunately not possible to tell for sure from
the available material whether Stupendemys had flippers or feet.
Other fossil pelomedusid genera include Polystemon with
three species from the Eocene to Oligocene of France, Paraliche-
lys with three species from the Oligocene and Pliocene of France
and Spain, Stereogenys with three species from the Eocene and
Oligocene of Egypt, Platycheloides with one species of probably
Cretaceous age from Malawi, Shweboemys with three species
from the Eocene to Pliocene of Egypt, Pakistan, and Burma, Car-
teremys with a single species from the early Tertiary of India, and
Elochelys with a single species from the Upper Cretaceous of
France. Other genera that have been named include Bantuchelys,
81
www.Ebook777.com
Dacochelys, Anthracochelys, and Calochelys, but these must re
main sub judice. The genus Stereogenys is very closely related to
Podocnemis; the species Stereogenys podocnemoides especially is
very similar to Podocnemis stromeri. However, Stereogenys is
characterized by an apparently constant separation of the anteri-
ormost neural from the nuchal bone. The entoplastron is small and
elongate; in S. podocnemoides the intergular separates both
gulars and humerals, while in S. cromeri the pectorals also are
partially separated. A decrease in size of all the anterior plastral
elements leads to S. libyca. The posterior plastral notch is fairly
large in this genus, and the skull is very short and broad A new
species, Stereogenys salamanticensis, has recently been de
scribed from the Eocene of Spain.
Platycheloides nyasae, of uncertain age from Malawi, is
thought by Zangerl to be a primitive relative of Pelomedusa. It is
characterized by the humero-pectoral sulcus lying well behind the
entoplastron, but the front and rear margins of the plastron are un
known. Shweboemys includes three species, S. antiqua from the
Eocene of Egypt, S. pilgrimi from the Pliocene or Pleistocene of
Burma, and S. gaffneyi from the Miocene of Pakistan. In this
genus the carapace was ridged, with the pleurals flat rather than
curved; the anterior plastral lobe was very short and semicircu
lar, and the skull, although Podocnemis-like in dorsal aspect, had
a broad secondary palate with a narrow median cleft such as is
found in Stereogenys. The carapace had a cordiform vertical pro
file, tapering to a point posteriorly. These were large turtles, since
the type skull of S. pilgrimi had an estimated basicranial length of
about five inches.
Carteremys leithi was a small species; the two best-preserved
shells available measure 7y3” and 8” respectively. The mid-plas-
tral area is not represented in the fossils, so it is not known if Car
teremys had mesoplastra and thus its status as a pelomedusid is in
some doubt. It was for many years considered to be a chelid, but
various characters, including the absence of a nuchal scute, the
very large intergular, the relative proportions of the first and
second vertebral scutes, the lack of a parieto-squamosal bar and
the presence of a jugo-quadratojugal bar, combine to suggest that
it does not belong in the Chelidae.
The modern genus Pelusios is known from fossils referred to
the species P. dewitzianus and P. blackenhomi from the Pliocene
and Miocene of Egypt respectively. Pliocene Pelusios fossils are
also known from the Baringo District of Kenya and from Omo in
Chad; the latter material appears to be referable to the modern
species P. sinuatus. An Oligocene fossil from Egypt has been con
sidered ancestral to the modern helmeted turtle, Pelomedusa sub-
rufa, and has been named Pelomedusa progaleata.
82
CRYPTODIRA
The Cryptodira are the dominant turtles in the world today,
and most of the known families of this suborder still have living
representatives. The essential definitions of the various families
thus were given earlier.
PLATYSTERNIDAE
This family is represented by a single species in south-east
Asia at the present time. Mlynarski considers Scutemys tecta,
from the Upper Cretaceous of China, to be referable to the Platy-
stemidae, although other authors have placed this same species in
either the Plesiochelyidae or the Dermatemyidae at different
times. Another form, Macrocephalochelys pontica from the Plio
cene of the Ukraine, is only known from an incomplete skull; since
the occipital region is lacking, it is not certain that this species
should not be referred to the Chelydrldae. A new species, Plant-
plastron tatarinovi, rather similar to the living Platystemon, has
recently been described from Oligocene formations of Russia.
CHELYDRIDAE
The snapping turtles are today represented by two purely New
World genera, Chelydra and Macroclemys. Eight fossil species of
Chelydra are recognized; the earlier ones (Oligocene to Pleisto
cene) are principally European, but American forms are repre
sented in the Pliocene to Recent periods. Macroclemys first
appears in the Miocene of North America; extinct forms of this
genus include M. schmidti from the Miocene of Nebraska and M.
auffenbergi from the Pliocene of northern Florida. Both are struc
turally very similar to the living M. temmincki. Other fossil chely-
drids include Acherontemys heckmanni from the Miocene of
Washington, presumably a small form since the 7” type specimen
lacked fontanelles in the carapace, and Hoplochelys, with seven
species from the early Tertiary of North America, which until re
cently were included in the Dermatemyidae. The earliest known
chelydrid is Gafsachelys from the Eocene of Europe and North
Africa. Chelydrops and Chelydropsis are two other genera repre
sented only by very fragmentary material and provisionally refer
red to the Chelydridae. Chelydropsis is known from ten species
from Europe, Asia, and possibly Africa.
KINOSTERNIDAE
The fossil record sheds little light on the origin of the genus
Kinosternon: one fossil species, K. arizonense, from the Pliocene
and Pleistocene of the south-western United States and northern
Mexico, has been described. It appears to be very similar to the
living K. flavescens, though it was slightly larger. The turtle Xeno-
chelys formosa, from the Oligocene of South Dakota, shows char-
83
www.Ebook777.com
acteristics of both the Dermatemyidae and the Staurotypinae; it
has the narrow shell and reduced, cruciform plastron of Stauro-
typus, but the smooth carapace and unhinged anterior plastral
lobe of Dermatemys. The axillary and inguinal scutes are in broad
contact on each side. Nine plastral scutes were present, in contrast
to the eight of Staurotypus and the eleven of Dermatemys (though
some fossil dermatemyids had other than eleven). A skull was
found with the type specimen of Xenochelys, but it is not certain if
the two specimens are definitely derived from the same animal.
The skull represents a more specialized turtle than does the shell.
DE RM ATE M YID AE
Although this family is today known only from the single spe
cies Dermatemys mawi, a number of fossil genera are known from
the Upper Jurassic onwards. The most clearly-defined and best-
known are North American, but different authors list various num
bers of rather doubtful genera from the Eocene to Miocene of Asia,
Europe, and Africa.
The New World dermatemyids had well-defined inframargi
nals, non-serrated carapace margins, and a tendency toward devi
ant numbers of plastral scutes. In some genera, including Derma
temys, there is a tendency for the dorsal scute boundaries to dis
appear. In several of the genera the posterior part of the neural
bone series is interrupted by pairs of contiguous pleurals. In the
genus Adocus, from the Upper Cretaceous and Paleocene of North
America, the posterior marginal scutes are greatly elongate,
while the plastron is rather small and has seven pairs of plastral
scutes, of which the second pair is rather widely separated. In the
species Adocus punctatus the mid-plastral seam is highly irregu
lar, forming a sinuous line that departs greatly from the midline.
In the genus Basilemys, from the Upper Cretaceous of Alberta, the
plastron is larger and the posterior marginals are very short. In
the species Basilemys sinuosa the mid-plastral seam is even more
sinuous than in Adocus punctatus, though this may be a function of
age. In Agomphus tardus, one of the eight species of the genus
Agomphus from the Upper Cretaceous and Early Tertiary of
North America, the carapace is steeply humped, the marginals
are very short, the plastron is rather small, and the scute bounda
ries of both carapace and plastron are greatly reduced. In the
genus Baptemys, from the Eocene of North America, the neural
series is uninterrupted but only five pairs of plastral scutes are
present.
The Old World genera of this family that have been reported
include Lindholmemys, from the Upper Cretaceous of Russia,
Sinochelys from the Lower Cretaceous of Asia, Trachyaspis from
the Eocene to Miocene of Europe and Africa, Heishanemys, Pei-
shanemys, and Tsaotanemys from the Cretaceous of China, Parta-
84
nemys from the Eocene of Europe, and Peltochelys from the

Upper Cretaceous of Belgium. Among the less well-known North
America genera are Homorophus, Zygoramma, Compsemys, No-
tomorpha, Anosteira, Pseudanosteira, Kallistira, and Alamo-
semys.
SINEMYDIDAE
This family, first described by Yeh Hsiang-k’uei in 1963, is
closely related to the Emydidae, but has the cross-shaped plastron
of more primitive cryptodires. Two species are recognized: Sine-
mys lens from the Lower Cretaceous of Shantung and Kansu,
China, and Manchurochelys manchuensis from the Upper Jurassic
of Liaoing, China. In the former species a mid-plastral fontanelle
is present; in the latter, there are two open fontanelles in each
bridge.
EMYDIDAE
This family is known from the Eocene up to the present day,
but the diversity of known fossil species is small compared to the
large number of living species and subspecies. One of the
most peculiar fossil genera is Clemmydopsis, with two species
from the Miocene and Pliocene of Germany and Hungary. In this
genus the plastron, the carapace margin, and the posterior
centrals and costals show the typical emydid arrangement, but the
first three centrals are laterally expanded so that they meet the
marginals and completely eliminate the first two pairs of costals.
Shell of Clem
mydopsis
sopronensis,
showing the wide
anterior central
scutes and only
two pairs of
costals. Redrawn
from Boda, 1927.
Another fossil emydid genus was Broilia, from the Oligocene

of France. Four species are recognized. These were large turtles,
up to about 32 inches in length, with elevated, elongated shells. The
plastron was not immoveably sutured to the carapace at the brid
ges, but rather was attached by cartilage. The entoplastron was
85
www.Ebook777.com
large and rounded and was not traversed by the humero-pectoral
sulcus. A deep notch was present between the xiphiplastra.
Three species of the genus Palaeochelys have been described
from the Oligocene of Mongolia and the Oligocene and Pliocene of
Europe. These turtles were much smaller than the species of
Broilia, and although sharing the elongate, elevated shells of tur
tles of that genus, usually had the humero-pectoral sulcus passing
through the entoplastron. A triangular interanal notch was pre
sent. This genus appears to be related to the living Geoemyda and
Ocadia.
In North America, the modem genera Terrapene and Pseu-
demys are known from a diversity of fossil material. An extinct
grassland box turtle, Terrapene omata long insulae, is known from
the Pliocene and early Pleistocene of Oklahoma and Kansas. A
subspecies, Terrapene Carolina putnami from the Pliocene and
Pleistocene of the southern and central United States (fossils
known from Florida, western Texas, and Kansas), was the largest
known box turtle, with a carapace about one foot in length. It had
the flared carapace margin that demonstrates close relationship
to Terrapene Carolina major, the largest living subspecies.
Fossil Pseudemys material from the United States demon
strates that the nelsoni-rubriventris line was already established
in Florida in the Miocene, and in Kansas and Idaho in the Pliocene.
Most of the known fossils of this genus, however, can be assigned
to the scripta group. Pseudemys platymarginata is from the
Irvingtonian (Pleistocene) of Florida and Texas. The extinct Pseu
demys scripta inflata was similar to certain specimens of the
modem P.s. scripta, but had exaggerated sculpturing, especially
of the nuchal area, as well as unusually sharp paired serrations on
the posterior peripheral bones. It is probable that the Pseudemys
line originated from a form similar to Echmatemys, from the
Eocene of North America.
PT Y CHOG ASTRID AE
This family, whose validity is not universally accepted, in
cludes the 27 or so species of Ptychogaster, a. genus known from
Eocene to Miocene deposits of Europe. The family in many ways
appears to bridge the gap between the Emydidae and the Testudi-
nidae, though the presence of a hinged plastron with moveable
hind lobe suggests a degree of specialization that would argue
against Ptychogaster being a direct ancestor of the Testudinidae.
TESTUDINIDAE
Fossil tortoises are known from many parts of the world from
the Eocene onwards. It is probable that the ancestral species were
not dissimilar to turtles of the living subgenus Manouria of the
genus Geochelone. One of the characteristic features of advanced
86
tortoises, the alternating octagonal and quadrilateral neural

bones, was not developed In such early forms as Stylemys nebras-
kensis, a possible ancestor of the genus Gopherus, from the Oligo
cene of the western United States. A number of other Oligocene
and Miocene species of this genus have been described, including
S. capax, S. calaverensis, S. conspecta, and S. oregonensis from
the New World, and S. botii, S. canetotiana, S. frizaciana, S. pyg-
mea, and S. karakolensis from Europe and Asia, some only known
from very fragmentary material. In this genus the pleural bones
showed very little alternation in width, a symphyseal dentary
groove was present, and a premaxillary ridge, similar to that of
Gopherus, was also present. At the present time, Gopherus is the
only tortoise genus in North America. The genus is known in the
fossil record from the Oligocene to the present; the widely dispar
ate ranges of the extant forms are a result of Pleistocene climatic
changes which exterminated the intermediate populations. About
twenty species have been described, some of them considerably
larger than those living today.
The still widespread genus Geochelone was long a dominant
member of the Nearctic fauna, as well as of the West Indian Is
lands, but in the New World the genus is now restricted to South
America and eastern Panama, and the Galapagos Islands. Popula
tions of Geochelone carbonaria on several of the West Indian Is
lands today are almost certainly introduced.
In the West Indies, fossil tortoises of large size are known from
Cuba (Geochelone cubensis), Sombrero (Geochelone sombreren-
sis), Nevassa (G. nevassae), and Mona (G. monensis); presum
ably many of the intermediate islands also once had tortoise popu
lations. These are Pleistocene and sub-Recent forms. The Mona
tortoise is referred to a separate subgenus (Monachelys) because
of its peculiarly elongate and narrow-waisted first dorsal verte
bra, as well as its shallow palate and feebly arched vomer.
In North America, one of the most interesting groups of tor
toises was the subgenus Hesperotestudo of the genus Geochelone.
This subgenus is characterized by the fusion of the ossicles and
scales dorsal to the tail, forming a large supracaudal ‘buckler,’
presumably for protection of the tail region. The subgenus presu
mably originated from the primitive Eocene subgenus Manouria
(which is still represented by two living Asiatic species). The ear
liest known species are amphithorax and brontops from the Lower
Oligocene. By the mid-Oligocene two distinct lines of evolution of
Hesperotestudo were apparent: a smooth thin-shelled line charac
terized by the Miocene species osbomiana, impensa, and augusti-
ceps, and the early Pliocene orthopygia, and a rugose, thick-shel
led line characterized by the early Pliocene pertenuis, the late
Pliocene turgida, and the Pleistocene equicomes, wilsoni, and in-
cisa.
87
www.Ebook777.com
Another North American subgenus was Caudochelys, in which
the limbs and tail were heavily armored with dermal ossicles, but
no consolidated supracaudal buckler was present. Relationships
within this subgenus remain somewhat obscure, beyond the fact
that the species showed a steady increase in size from the Upper
Eocene to the Pleistocene. The earliest species possibly referable
to Caudochelys is uintensis, from the Upper Eocene of Utah. Other
species are G. ligonia from the Lower Oligocene of Colorado; G.
arenivaga from the Miocene of Nebraska; G. tedwhitei from the
Lower Miocene of Florida, G. hayi from the Pliocene of Florida, G.
rexroadensis from the Pliocene of Kansas, G. laticaudata from the
Pleistocene of the Western United States, and G. crassiscutata
from the Pleistocene of Florida. The latter was an enormous beast,
with a thick carapace up to four feet or more in length. G. hayi, of
which G. louisekressmani is probably a synonym, was also very
large. There are numerous other fossil Geochelone species from
the Oligocene, Miocene, and Pliocene of North America, but all are
presently known from such incomplete material that subgeneric
designations cannot be made.
The other fossil tortoise genera are less well known. They in
clude Floridemys, Cymatholcus, Achilemys, Cheirogaster, Colos-
soemys, and Sinohadrianus. Floridemys nanus is known only from
shell material from the Florida Pliocene. It was a very small spe
cies characterized by the transverse gulohumeral sulcus. Cyma
tholcus, represented by the two species C. longus and C. schucher-
ti, is a poorly known Eocene genus that may be related to Stylemys
or ancestral to Hesperotestudo. It has recently been suggested by
Auffenberg that Cymatholcus should be considered a subgenus of
Geochelone; it is distinguished from the related Manouria by the
narrower shell and narrow pectoral scutes. Achilemys is the gene
ric name given by Hay to certain fragments that are so incomplete
that they probably do not even deserve taxonomic recognition.
Cheirogaster is founded on a perfect shell from the Eocene of
Europe. It is reported to lack the entoplastron—certainly a charac
ter worthy of generic recognition, but unfortunately the describer,
Bergounioux, was wrong in practically everything he ever wrote,
so it is important that the type be re-examined by a competent
palaeontologist. Most of the material referred by Rodrigues to
Colossoemys is not even chelonian, except for an indeterminate
plastral fragment. Another genus, Sinohadrianus from the Eocene
of China, appears to be a primitive tortoise, but the single imper
fect shell known shows almost no distinctive characters.
In tropical America, a large species, Geochelone costaricen-
sis, is known from an uncertain period (possibly Oligocene or Mio
cene) of Central America. In South America, Geochelone gringo-
rum from the Miocene of Patagonia is very close to the living G.
88
chilensis. The newly-described species G. hestema, from the late

Miocene of Colombia, appears to be intermediate in many respects
between the living G. carbonaria and G. denticulata, and may be
ancestral to both.
In the Old World, numerous fossil tortoises of large size are
known, typified by the Eocene Geochelone (Asterochelys) ammon
from Egypt. The largest known tortoise ever was Geochelone
(Megalochelys) sivalensis (also known as Colossochelys atlas)
from the Pleistocene of the Sivalik Hills of India, and also recorded
from the Indonesian islands of Timor, Java, and Celebes. The
carapace length of this species, judging by the nearly complete
specimen in the American Museum of Natural History and the
isolated bones and shell fragments in the British Museum and
Leiden Museum, was approximately six feet. The reconstructed
shell on exhibit in the British Museum, which is over eight feet
long, is exaggerated. A somewhat smaller species, Geochelone
(Megalochelys) gadowi, also with strongly forked gulars, is known
from the Pleistocene of Mauritius.
The genus Testudo is known from a number of fossil species
from Europe, Russia, and the Middle East. The paleontologist
Chkikvadze has recently proposed a new genus, Protestudo, for
certain of the more primitive species customarily assigned to the
genus Testudo, and he includes the living species hermannl in this
new genus. Protestudo is thus known from the Miocene to Recent
of Europe, and from the Oligocene to Pliocene of Asia. The species
assigned to Protestudo include antiqua, aralensis, bessarabica,
hermanni, hipparionum, honanensis, hypercostata, Jcalksburgen-
sis, kegenica, oskarkuhni, promarginata, schansiensis, schensien-
sis, sphaerica, szalai, tunhuanensis, turgaica, and yushensis.
Chkikvadze considers Protestudo to be ancestral to the extinct
genus Agrionemys as well as to Testudo. Chkikvadze also consid
ers his genus Ergilemys, from the Oligocene of Mongolia, to be
ancestral to Protestudo, Manouria, Geochelone, and Indotestudo,
but it would seem that the Oligocene would have been too late for
the common ancestor of these ancient genera and subgenera.
TRIONYCHIDAE
Despite their specialized form, the soft-shelled turtles (Triony-
chidae) are one of the most ancient of living families, having a fos
sil record extending at least as far back as the Cretaceous. Even
fragmentary remains can usually be assigned to this family since
the bones of both carapace and plastron are usually heavily pitted
in a characteristic fashion. Perhaps the most ancient form is Sin-
aspideretes wimani, described by Young and Chow in 1953, which
may be of Upper Jurassic age. This species had a relatively deep,
keeled carapace, lacking the peripheral bones, and with a thick,
89
www.Ebook777.com
broad, smooth plastron lacking the usual callosities. Traces of
scute boundaries are discernible on the carapace. Better known is
the genus Plastomenus, known from about eleven species from the
Cretaceous and Eocene of North America, with a single newly-
discovered form (P. mlynarskii) from the Eocene of eastern Ka
zakhstan. In this genus the plastral bones were large and were al
most or completely contiguous. Generally the entoplastron was
absent and the epiplastra were fused, as in modem soft-shells, but
preplastra were absent. However, in the type specimen of Plasto
menus tentillus Hay, the composite epiplastron was absent, though
this part may have been lost before fossilization.
Several of the modern genera have fossil histories. Fragments
of Trionyx are exceedingly plentiful in many fossil beds, and not
fewer than 180 species have been described. These are known from
the Eocene to Pliocene of Europe, from the Eocene to Recent of
North America, from the Oligocene to Recent of Asia, and from the
Pleistocene to Recent of Indonesia. It is customarily stated that
the Trionychidae have never inhabited South America, but Roger
Wood recently showed me a large fossil Trionyx (or at least triony-
chid) hyoplastron which he had collected in Venezuela.
The modern African genus Cycloderma is known from one fos
sil species (C. victoriae) from the Miocene of East Africa. Three
Pliocene species of Lissemys have been described from the Sivalik
Hills of India. Two Pleistocene species of Chitra from Java are
known, while certain possibly trionychid fossil fragments from
northern Australia may be referable to Pelochelys.
CARETTOCHELYIDAE
The single living species of this family is characterized by the
overall sculpturing of the superficial bones of both the shell and
the skull and by the absence of scutes, although rudimentary cen
tral scute boundaries are discernible in juveniles. Although now
confined to the southern part of New Guinea and to one or two
rivers in northern Australia, the family was once very widely dis
tributed, being known from the Paleocene to the Oligocene of
Europe, and from the Eocene of North America and Asia. The liv
ing genus Carettochelys is known from the Miocene onwards. It is
usually placed in a separate subfamily (Carettochelyinae), distin
guished from the extinct Anosteirinae in which the plastron is
rather narrow and cruciform and in which sulci, at least on the
carapace, are often well-developed.
In the genus Anosteira, of which 19 species from the Eocene
and Oligocene of eastern Asia, Europe, and North America are
known, the entoplastron was triangular as in the living Carettoche
lys, but the hind plastral lobe was as narrow as that of Stauro-
typus, and most of the carapacial sulci were apparent. In Ano
steira ornata from the Eocene of Wyoming, for example, the costal
90
and marginal scutes were defined by almost complete sulci, but

the anterior central was longitudinally split, and the space normal
ly occupied by the second, third, and fourth centrals was occupied
by a single elongate scute. In Pseudanosteira five centrals were
present, but they were of very unequal size, the fourth being ex
tremely small, while the first and third had projecting rami that
almost enclosed the second.
Sulci are absent in the genera Allaeochelys and Akrochelys
from the Eocene of France. In Allaeochelys the peripheral bones
usually did not form a complete series and were tightly sutured to
the costals. The bridge was bony and rigid. In Akrochelys the cara
pace was well ossified and had seven neural bones. The peripheral
bones formed a complete series.
CHELYCARAPOOKIDAE
This family was erected recently to accommodate a single
species, Chelycarapookus arcuatus, known from a single fossil col
lected at Carapook, Victoria, Australia, in 1919. The specimen was
for years considered to be referable to the modem side-neck spe
cies Emydura macquarri. The affinities of the new family, if in
deed it is valid, have yet to be determined.
NANHSIUNGCHELYIDAE
This family includes the single species Nanhsiungchelys wuch-
ingensis, described in 1965 by Yeh Hsiang-k’uei. The holotype,
which includes carapace, plastron, skull, lower jaw, and various
other bones, is from the Upper Cretaceous of Kwangtung, China.
The family may be most closely related to the Carettochelyidae;
the carapace, 970 mm long, has a sculptured surface, distinct scute
sulci, and an exceedingly strong nuchal emargination. The neural
bones are somewhat indistinct, but at least one octagonal neural is
visible. The skull is 185 mm wide, and is of a most peculiar shape,
with very widely separated eyes, exceedingly expanded nasal
opening, and a highly concave anterior profile.
The plastron of Nanhsiungchelys, like the carapace, is strong
ly sculptured; the bridge is wide and the entoplastron very large.
TOXOCHELIDAE
This family of extinct sea turtles is probably derived from the
amphichelid family Thalassemyidae, and it in turn is closely rela
ted to the modern Cheloniidae. It will thus be seen that the Crypto-
dira, as presently defined to include the sea turtles, are at least bi-
phyletically derived from the ancestral Amphichelydia. It would
thus seem that the present allocation of the sea turtles to the Cryp-
todira (which, after all, derive their name from the retractile
nature of the extremities, a manifest impossibility in the sea tur
tles ) may be incorrect. A new classification of the turtle families
91
www.Ebook777.com
by Rainer Zangerl, given later in this chapter, goes some way to
ward correcting this problem.
The Toxochelidae varied in their degree of aquatic specializa
tion, some probably being near-shore dwellers while others may
have been pelagic. The skull roof was slightly emarginate from be
hind, and the skull sometimes had a secondary palate. The plas
tron was cruciform, with the distance between the axillary and in
guinal notches not more than 60% of the half-width of the plastron.
The carapace and plastron almost always showed lateral fontanel-
les. The forelimbs were developed into flippers, but the hind feet
were more similar to those of snapping turtles. The carapace was
usually oval or circular, and two suprapygal bones were present.
The Toxochelidae were a Late Cretaceous family whose fossils
have been found mainly in North America.* Three sub
families are recognized: the Toxochelyinae is the more primitive
and is characterized by the absence of a secondary palate, the an
teriorly narrow triturating surfaces, the small nasal bones, the cir
cular (rarely cordiform) carapace, the non-serrated shell margin,
the small central and lateral plastral fontanelles, and the flat neu
ral bones. The genus Toxochelys itself is known from the species
latiremis, moorevillensis, barberi, atlantica, weeksi, and broumi.
The species are distinguished largely by differences in the shape of
the plastron. The carapace is oval or nearly circular, with mode
rate or extensive lateral fontanelles, and also a pair of post-nuchal
fontanelles. The related species Thinochelys lapisossea, from the
Selma Formation of Alabama, is placed in a separate genus be
cause the lateral fontanelles are very small, the postnuchal fonta
nelles are absent, and the neural bones meet each other at very
strongly curved sutures. There are also differences in plastral pro
portions and in the degree of serration of the inner edge of the hyo-
and hypoplastra. The third genus, Porthochelys, with the single
species P. laticeps, from the Niobrara Formation of Kansas, had a
wide, circular carapace, with very small lateral fontanelles and a
relatively short, broad, and massive skull. Only one specimen is
known, and it is possible that the species did not live in the Nio
brara Sea, but rather in adjacent coastal fresh water, so that fos-
silization in the chalk would have been a rare event.
In the second subfamily, the Osteopyginae, a well-developed
secondary palate was always present and nasal bones were prob
ably absent. The neural bones were unkeeled, and the carapace
margin was at most only slightly serrated. The mandibular sym
physis was broad—probably more than one third the length of the
ramus. A deep pit was present in the postero-medial face of the
second peripheral bones for reception of the anterio-lateral prong
* European and African toxochelids are now known.
92
of the hyoplastron. Only a single species, Osteopygis emarginatus,

is currently recognized, though this species is sufficiently variable
for no fewer than thirteen synonyms to have been perpetrated in
the literature. Eight of these were described as species of Osteo
pygis, but others were placed in the genera Catapleura, Propleura,
Lytoloma, and Erquelinnesia. The skull of Osteopygis is so similar
to that of the cheloniid sea turtles that the genus would probably
have been placed in that family were shell material not available.
The carapace shows great variability in the presence or absence of
lateral fontanelles.
The species Rhetechelys platyops, based on a large, incom
plete skull from the late Cretaceous of New Jersey, may also be
referable to the Osteopyginae; it differs from Osteopygis not only
in its larger size, but also in the presence of a deep pit in the palate
behind the beak which suggests the presence of a strongly hooked
lower jaw. Also, the snout is relatively slender and pointed.
The most specialized toxochelids are included in the subfamily
Lophochelyinae. Broad triturating surfaces were present, and the
beginnings of a secondary palate are evident. Small nasal bones
were present. The carapace was oval to cordiform, and a serrated
median keel, sometimes with distinct epineural bones on the tips of
the tubercles, was present. Large fontanelles were always present
in the carapace and plastron. Three genera are known—Lopho-
chelys, Ctenochelys, and Prionochelys. Three species of Lopho-
chelys have been described—natatrix and niobrarae from the Nio
brara of Kansas, and venatrix from the Mooreville Chalk of Ala
bama. These deposits were both of Upper Cretaceous age. The
skull of this genus is unknown, but the neural keel lacked the sepa
rate epineural bones present in the other genera of this subfamily.
Extensive carapacial fontanelles, including a postnuchal pair,
were present. The carapace edge was moderately serrate. In
young specimens (at least) the pygal and suprapygal bones were
not in contact.
The genus Ctenochelys is represented by four species—two
(tenuitesta and acris) from the Mooreville Chalk, and two (steno-
pora and procax) from the Niobrara. The skull of Ctenochelys is
fairly massive, with very heavy, strong jaws. The carapace, on the
other hand, was composed of very thin, spongy bones. At least
three of the tubercles of the dorsal keel were tipped with separate
epineural bones. Postnuchal fontanelles were present, and the
carapace margin was moderately to deeply serrate. In adults the
pygal and suprapygal bones had a tenuous connection. Wide
lateral fontanelles were present in the carapace. The shell was
typically strikingly elongate.
Prionochelys, with the three species nauta, matutina, and
galeotergum, was the most specialized genus of the Toxochelidae.
93
www.Ebook777.com
These must have been striking-looking turtles, with highly serra
ted carapace margins and very strong median tubercles, each
capped with an epineural bone. The overall shape of the shell was
cordiform, with a pronounced excavation of the anterior nuchal
rim. There were no postnuchal fontanelles in adults. The plastron
is poorly known, but the xiphiplastra are reportedly similar to
those of Ctenochelys. The skull also is only known from fragments,
but these suggest that the head was large, and the mandible mas
sive. The palatal region showed the typical lophochelyine incipient
secondary palate, with the beginnings of an undershelving of the
internal nasal openings.
CHELONIIDAE
The modem sea turtles of the family Cheloniidae are clearly
fairly closely related to the Toxochelidae, the principal differences
being the large plastron and secondary palate in the Cheloniidae,
and the cruciform plastron and usually absent secondary palate of
the Toxochelidae. In many ways the Cretaceous genus Catapleura
and the living Lepidochelys can be considered to bridge the two
families.
Tracing the ancestry of the Cheloniidae is complicated by the
fact that the Cretaceous fossils assignable to this family tend to be
considerably more specialized than both the early Tertiary fossils
and the living genera. Thus we have little material representing
the main line of cheloniid evolution and are instead restricted to
glimpses of some of the early dead-end evolutionary branches. The
difficulties of establishing a natural division of the living cheloniid
genera into subfamilies are discussed later. However, if we accept
the controversial notion that Chelonia and Eretmochelys may be
placed in one subfamily, the Chelonini, and Caretta and Lepido
chelys in another, the Carettini, it is possible to identify certain
fossil genera as Chelonini ancestors or relatives, while certain
others can be associated with the Carettini.
Pliocene fossil cheloniids known at the present time include
Chelonia sismondai, a close relative of the living Chelonia mydas,
and Pachychelys robusta, a specialized, solidly-built form known
from a skull fragment, the lower jaw, and the pleural bones. A
third Pliocene species, “Chelonia” gastaldi, is now known to be a
dermochelyid. Several Miocene cheloniids are known; these include
Chelonia girundica, which was apparently a gigantic relative of
Chelonia mydas, and Chelonia califomiensis, which may be a syn
onym of girundica—it is only known from a single femur; Euclas-
tes melii, which, having divided neural bones, may be a Carettine
ancestor, though it lacks the small extra costal scute of modem
Carettini; Peritresius and Syllomus are specialized genera with
bony ornamentation on the skull; they are distinguished by the
94
presence in only the former of intercostal fontanelles. Procolpo-

chelys grandaeva, from the Miocene of New Jersey, had the divi
ded neurals that suggest Carettine affinities; the carapace was
elongate, with strong sulci indicating only four pairs of costal
scutes and with large lateral fontanelles. Kurobechelys tricarinata
was a cheloniid from the Miocene of eastern Asia.
Oligocene cheloniids include “Chelonia” gwinneri, which from
its bridge index may be more properly associated with the Caret
tini, and Glarichelys knorri, a European form distinguished by its
elongate snout, short mandibular symphysis, and very narrow
interorbital bridge. Mlynarski describes this as a very small turtle
(about 20 cm. long) with very long flippers; but it is more likely
that the material available to Mlynarski was simply derived from
very young turtles.
Less well known Oligocene species include Chelonia (or Bryo-
chelys) waterkeyni, Chelonia vanbenedeni, Chelyopsis holstati-
cus, Chelyopsis littoreus, and Carolinochelys wilsoni. The last-
named, only known from a skull from the Oligocene of North Caro
lina, may be a Lepidochelys ancestor, though the presence of a
specialized pattern of ridges on the palate would tend to argue
against it being in the direct line of ancestry. It is even possible
that the skull that has been labeled Carolinochelys could corres
pond to the skull-less material on which Procolpochelys has been
based.
Many Eocene cheloniids have been described, but few are well
known. Those forms such as Eochelone, Erquelinnesia, Lytoloma,
and Argillochelys may just as reasonably be placed in the Toxo
chelidae as in the Cheloniidae. Several North African Eocene che
loniids, including Gafsachelys phosphatica, Thalassochelys phos-
phatica, and Euclastes douvillei, have been described; the last two
should probably be placed in other genera.
Cheloniids were apparently world-wide even as far back as the
Cretaceous, but most of the genera recorded from that era could
just as well be placed in one of the other sea turtle families.
PROTOSTEGIDAE
This family of extinct sea turtles is only known from Upper
Cretaceous deposits of the United States; * included are some very
large species, culminating in Archelon ischyros, one of the
largest turtles that ever lived. The family is characterized by a
number of features, which include the long hooked beak, the highly
fenestrated carapace (in the larger genera, the carapace struc
ture was similar to that of most hatchling turtles, the pleural bones
only touching at their proximal ends), the roofed skull, the absence
* Cimochelys benstedi, from the Upper Cretaceous of England,

may be a protostegid.
95
www.Ebook777.com
of a secondary palate, and the reduced xiphiplastra. Nasal hones
were present, the plastron was composed of heavy plates (though
always with a median fontanelle), the neck was short and presum
ably non-re tractile, epiplastra were sometimes absent, and there
were often as many as nine ribs/pleurals.
Carapace and plastron of a gigantic extinct protostegid turtle, Archelon

ischyros. Redrawn from Suchanow, 1964.
The more primitive protostegids of the genera Chelosphargis

and Calcarichelys are placed in a separate subfamily, the Chelos-
pharginae. They were of relatively small size, with relatively well-
ossified shells. Chelosphargis advena was the more primitive of
the two species; it had a blunt, very low median keel, an even cara
pace margin, and a pygal bone completely separating the eleventh
pair of pleurals. Calcarichelys gemma, also known from both the
Niobrara Chalk of Kansas and the Mooreville Chalk of Alabama,
was a spectacular form with a very wavy carapace margin and
with every other neural bone showing a high, triangular lateral
profile, so that a uniquely serrate median keel was formed. Late
ral fontanelles were moderate, and the pygal bone was very nar
row.
The subfamily Protosteginae contained two genera, Protoste-
ga and Archelon. Protostega, with four species from Kansas,
Texas, and Alabama, had a skull nearly as wide as long and a
wavy dorsal keel. Large hyo- and hypoplastra were present, but
the xiphiplastra were small and the epiplastra lacking. The cara-
pacial fontanelles were so large that the costal plates were essenti
ally vestigial. Of the two species of Archelon, A. copei was the
smaller and A. ischyros the larger—the latter reached a total
96
The giant extinct Upper Cretaceous sea turtle Protostega gigas, photograph
ed at the Carnegie Museum, Pittsburg.
97
www.Ebook777.com
length (including the very large skull) of about ten feet. The genus
is distinguished by the elongate head with downcurved beak and
the even neural keel. Contrary to earlier belief, it is now known
that all of the protostegids had dorsal scutes. The best-known spe
cimen of Archelon ischyros is a well-mounted example in the Yale
University Museum, complete except for one of the hind feet which
was probably severed by a predator during the life of the animal.
DERMOCHELYIDAE
The highly divergent family Dermochelyidae is discussed in
some detail later, in the account of the single living species, Der-
mochelys coriacea. The origins of the family are uncertain, but it is
presumed that it arose as an early offshoot of the primitive
cheloniids. The Eocene dermochelyid genus Psephophorus is
known from about eight Eocene to Miocene species from North
Africa, Europe, and North America. None of the material is well-
preserved, but it is guessed that the typical adult length was be
tween three and four feet. Eosphargis gigas was another Eocene
dermochelyid, with two species known from England, Denmark,
and Belgium. It too was smaller than the living Dermochelys. The
genus Dermochelys itself is known from the Miocene onwards.
CLASSIFICATIONS
The classification of the living and extinct turtle families is by
no means unanimous; different authors who have emphasized
different characters have arrived at very divergent systems. The
six systems given below are all by modern authorities who have
considerable experience with both the living and the fossil groups.
The systems are the work of Mlynarski (1969), Chkhikvadze
(1970), Romer (1956), Williams (1950), Zangerl (1969), and
Gaffney (1975) respectively. All are commended to the reader’s
attention, and no attempt is made to suggest that one is ‘better’
than the others. Gaffney’s system is noteworthy in that it abandons
the concept of the Suborder Amphichelydia, places great emphasis
on the distinctness of the Triassic turtles of the family
Proganochelyidae, and places the kinosternids and the
dermatemyids together with the carettochelyids and the
trionychids in the superfamily Trionychoidea. These four families
are placed together on the basis of the presence of a caudifibularis
muscle, the reduced or absent stapedial artery, and seven skull
characters. Gaffney’s classification recognizes fewer families
than any of the others, but some taxa (such as the Jurassic
Solnhofia and the meiolaniids) are omitted because their
relationships are still uncertain.
98
(I) Mlynarski, 1969

Order: Testudines
Suborder: Amphichelydia
Family: Archaeochelydiidae (incerta)
Superfamily: Proganochelyoidea
Families: Proganochelyidae, Proterochersidae
Superfamily: Pleurosternoidea
Families: Pleurosternidae (Pleurosteminae,
Desmemydinae), Kallokibotiidae,
Plesiochelyidae, Thalassemydidae,
Aperotemporalidae (incerta)
Superfamily: Baenoidea
Families: Baenidae, Eubaenidae, Macrobaenidae,
Neurankylidae, Meiolaniidae
Suborder: Cryptodira
Superfamily: Chelydroidea
Families: Dermatemyidae, Sinemydidae, Chelydridae,
Kinostemidae (Kinosteminae,
Staurotypinae)
Superfamily: Testudinoidea
Families: Emydidae (Batagurinae, Emydinae),
Platystemidae, Ptychogastridae,
Testudinidae
Superfamily: Chelonioidea
Families: Toxochelyidae (Toxochelyinae,
Lophochelyinae, Osteopyginae),
Protostegidae (Chelospharginae,
Protosteginae), Desmatochelyidae,
Cheloniidae
Superfamily: Dermochelyoidea
Family: Dermochelyidae
Superfamily: Trionychoidea
Families: Carettochelyidae (Anosteirinae,
Carettochelyinae), Trionychidae
(Cyclanorbinae, Trionychinae)
Suborder: Pleurodira
Families: Pelomedusidae, Eusarkiidae, Chelidae.
(II) Chkhikvadze, 1970
Order: Testudinata
Infraorder: Amphiplastra
Families: Proganochelyidae, Proterochersidae
Families: Baenidae, Eubaenidae, Kallokibotiidae
Infraorder: Pseudochelonoplastra
99
www.Ebook777.com
Families: Desmemydidae, Meiolaniidae (?),
(?) Neurankylidae, (?) Helochelydridae
Superfamily: Pleurostemoidea
Families: Pleurostemidae, (?) Platychelyidae
Superfamily: Pelomedusoidea
Families: Pelomedusidae (Bothrememydinae,
Eusarkiinae, Pelusiinae, Podocnemiinae,
Pelomedusinae), Chelidae (Hydraspinae,
Chelinae)
Infraorder: Chelonoplastra
Superfamily: Thalassemydoidea
Families: Thalassemydidae, (?) Macrobaenidae
Families: Cheloniidae, Protostegidae,
Desmatochelyidae
Superfamily: Dermochelioidea
Infraorder: Cruciplastra
Superfamily: Chelydroidea
Families: Sinemydidae, Toxochelyidae (Osteopyginae,
Toxochelyinae, Lophochelyinae),
Chelydridae
Superfamily: Carettochelyoidea
Families: Staurotypidae, Kinostemidae, Anosteiridae,
Carettochelyidae
Infraorder: Trionychoplastra
Families: Sinaspideretidae, Cyclanorbidae
(Lissemydinae, Cyclanorbinae),
Plastomenidae (Plastomeninae, Chitrinae),
Trionychidae
Infraorder: Planiplastra
Superfamily: Plesiochelyoidea
Families: Sinochelyidae, Plesiochelyidae, Adocidae,
Dermatemyidae
Superfamily: Platystemoidea
Family: Platystemidae
Families: Lindholmemydidae, Bataguridae,
Geoemydidae (Ptychogasterinae,
Geoemydinae, Sakyinae), Emydidae,
Testudinidae
100
(III) Romer, 1956

Order: Chelonia
Family: Proganochelyidae
Superfamily: Pleurostemidae
Families: Pleurostemidae (Desmemydinae,
Kallokibotinae, Pleurosteminae),
Plesiochelyidae, Thalassemydidae,
Aperotemporalidae
Families: Neurankylidae, Baenidae, Meiolanidae,
Eubaenidae
Families: Dermatemydidae, Chelydridae (Chelydrinae,
Staurotypinae, Kinosteminae), Testudinidae
(Emydinae, Testudininae, Platysteminae)
Families: Toxochelyidae (Toxochelyinae,
Lophochelyinae, Osteopyginae),
Protostegidae (Chelospharginae,
Protosteginae), Desmatochelyidae,
Cheloniidae
Family: Carettochelyidae (Carettochelyinae,
Anosteirinae)
Family: Trionychidae
Families: Pelomedusidae, Chelidae
Suborder: Eunotosauria
Family: Eunotosauridae
(IV) Williams, 1950
Order: Testudinata
Superfamily: Triassochelyoidea
Family: Triassochelyidae (Triassochelyinae,
Proterochersinae)
Superfamily: Pleurostemoidea
Families: Pleurostemidae (Pleurosteminae,
Desmemydinae, Kallokibotinae),
Plesiochelyidae, Thalassemydidae
101
www.Ebook777.com
Families: Baenidae (Baeninae, Meiolaniinae),
Neurankylidae, Eubaenidae.
Families: Dermatemydidae, Chelydridae (Chelydrinae,
Staurotypinae, Kinosteminae), Testudinidae
(Platysteminae, Emydinae, Testudininae)
Families: Toxochelidae (Toxochelyinae, Lytolominae),
Protostegidae, Cheloniidae
Family: Carettochelyidae (Carettochelyinae,
Anosteirinae)
Family: Trionychidae (Plastomeninae, Trionychinae,
Lissemydinae)
Family: Pelomedusidae (Bothremydinae,
Pleomedusinae), Chelidae
(V) Zangerl, 1969
Order: Testudines ( = Chelonia)
Families: Proganochelyidae, Proterochersidae,
Kallokibotiidae
Suborder: Mesochelydia
Families: Pleurostemidae, Neurankylidae, Baenidae,
Compsemyidae, Eubaenidae, Meiolaniidae,
Pelomedusidae, Chelyidae
Suborder: Metachelydia
Families: Plesiochelyidae, Thalassemydidae,
Sinemydidae, Aperotemporalidae,
Dermatemydidae, Chelydridae,
Toxochelyidae, Protostegidae, Cheloniidae,
Dermochelyidae, Carettochelyidae,
Trionychidae
Suborder: Neochelydia
Family: Testudinidae
(VI) Gaffney, 1975

Order Testudines Linnaeus, 1758
Suborder Proganocheiydia Romer, 1966
Family Proganochelyidae (Triassic)
Suborder Casichelydia Gaffney 1975a
102
Infraorder Pleurodira (Cope, 1868)

Family Pelomedusidae (Cretaceous-Recent)
Family Chelidae (Eocene-Recent)
Infraorder Cryptodira (Cope, 1868)
Parvorder Paracryptodira, Gaffney 1975b
Superfamily Baenoidea Williams, 1950
Family Glyptopsidae (Jurassic)
Family Baenidae (Cretaceous-Eocene)
Parvorder Eucryptodira Gaffney, 1975a
Superfamily Trionychoidea Gray, 1870
Family Kinostemidae (Oligocene-Recent)
Family Dermatemyidae (Cretaceous-Recent)
Family Carettochelyidae (Eocene-Recent)
Family Trionychidae (Cretaceous-Recent)
Superfamily Chelonioidea Baur, 1893
Family Plesiochelyidae (Jurassic)
Family Protostegidae (Cretaceous)
Family Toxochelyidae (Cretaceous-Eocene)
Family Dermochelyidae (Eocene-Recent)
Family Cheloniidae (Cretaceous-Recent)
Superfamily Testudinoidea Baur, 1893
Family Chelydridae (Paleocene-Recent)
Family Emydidae (Paleocene-Recent)
Family Testudinidae (Eocene-Recent)
BACKGROUND READING
Auffenberg, W. 1958. Fossil turtles of the genus Terrapene in Flori
da. Bull. Florida State Mus. 3 (2): 53-92.
__________ 1962a. A new species of Geochelone from the Pleisto
cene of Texas. Copeia 1962 (3): 627-636.
__________ 1962b. A redescription of Testudo hexagonota. Herpeto-
logica, 18 (1): 25-34.
__________ 1963. Fossil testudinine turtles of Florida genera Geo
chelone and Floridemys. Bull. Florida State Mus. 7 (2): 53-97.
__________ 1964. A redefinition of the fossil turtoise genus Stylemys
Leidy. J. Palaeontology 38 ( 2): 316-324.
__________ 1964b. A new fossil tortoise from the Texas Miocene,
with remarks on the probable geologic history of tortoises in
eastern U.S. Pearce-Sellards Series, Texas Memorial Mus.,
Univ. Texas (3): 1-10.
__________ 1966. A new species of Pliocene tortoise genus Geoche
lone from Florida. J. Palaeontology 40 (4): 877-882.
103
www.Ebook777.com
__________ 1971. A new fossil tortoise, with remarks on the origin of
South American Testudinines. Copeia 1971 (1): 106-117.
__________ 1974. Checklist of fossil land tortoises (Testudinae).
Bull. Florida State Mus., 18(3): 121-251.
Baird, D. 1964. A fossil sea-turtle from New Jersey. N.J. State
Mus. Inves., No. 1, pp. 3-26.
Bergounioux, F.M. 1938. Archaeochelys Pougeti nov. gen., nov.
sp., tortue fossile de Permien de l’Aveyron. Bull. Soc. geol.
France 5 (8), pp. 67-72.
_________ . 1952. Les cheloniens fossiles de Gafsa (Appendice).
Not. Mem. Serv. geol. Maroc 92, pp. 377-396.
__________ 1955. Chelonia. In: Traite de Paleontologie 5, pp. 487-
544.
Brattstrom, B.H. 1961. Some new fossil tortoises from western
North America with remarks on the zoogeography and palaeo-
ecology of tortoises. Joum. Palaeont., 35 ( 3): 543-560.
Brown, B. 1931. The largest known land tortoise. Nat. Hist., 31:
183-187.
Chkhikvadze, V.M. 1970a. On the origin of the modern Palaearctic
land tortoises. Bull. Acad. Sci. Georgian S.S.r. 57(1): 245-247.
__________ 1970b. Klassifikatsiya Podklassa Testudinat. 16th Annual
Session, Inst. Paleobiology. 10-11 Dec. 1970. Tesis Dokladov: 7-8.
__________ 1971a. On the history of the tortoise family Chelydridae.
Bull. Acad. Sci. Georgian S.S.R. 61 (1): 237-240.
__________. 1971b. New tortoise species of the Oligocene of
Kazakhstan and the systematic position of some of the fossil
species of Mongolia. Bull. Acad. Sci. Georgian S.S.R. 62 (2):
489-492.
__________. 1971c. Three new fossil species of land tortoises from
Zaisan Basin (Eastern Kazakhstan). Bull. Acad. Sci. Georgian
5.5. R. 64 (1): 245-258.
__________ 1971d. Pervaya Nachodka Tretichnoi Cherepaki Cemeist-
va Platystemidae. Paleontologichesi J., Moscow, 4: 137-139.
__________ 1972. On the systematic position of Tertiary gigantic
land tortoises of the Palearctic. Bull. Acad. Sci. Georgian
5.5. R. 65 (3): 745-748.
__________. 1973a. New Data on Tertiary turtles from Georgia.
Bull. Acad. Sci. Georgian S.S.R. 69 (3): 745-748.
__________ 1973b. Interrelations of forms and functions of the tur
tle shell showing the direction of natural selection. Acad. Sci.
Georgian S.S.R., Palaeontological Institute. VI: 65-78.
Collins, R.L. and W.G. Lynn. 1936. Fossil turtles from Maryland.
Proc. Amer. Philos. Soc., 76 (2): 151-174.
Collins, J.I. 1970. The chelonian Rhinochelys Seeley from the
Upper Cretaceous of England and France. Palaeontology, 13
(3): 355-378.
104
Dacque, E. 1912. Die fossilen Schildkroten Aegyptens. Geol. Palae-

ont. Abhandl., 14, pp. 275-337.
Gaffney, E.S. 1972. The systematics of the North American family
Baenidae (Reptilia, Cryptodira). Bull. Amer. Mus. Nat. Hist.,
147(5): 241-320.
__________ 1975. A phylogeny and classification of the higher cate
gories of turtles. Bull. Amer. Mus. Nat. His., 155(5): 387-436.
Gaffney, E.S. and R. Zangerl. 1968. A revision of the chelonian
genus Bothremys (Pleurodira: Pelomedusidae). Fieldiana,
Geol. 16 (7): 193-239.
Gilmore, C.W. 1931. Fossil turtles of Mongolia. Bull. Amer. Mus.
Nat. Hist. 59, pp. 213-257.
Glaessner, M. 1942. The occurrence of the New Guinea turtle
(Carettochelys) in the Miocene of Papua. Rec. Austral. Mus.
21 (2): 106-109.
Hay, O.P. 1908. The fossil turtles of North America. Carnegie Inst.
Washington, Pub. No. 75, pp. 1-568.
Hooijes, D.A. 1971. A giant land tortoise, Geochelone atlas (Fal
coner & Cautley), from the Pliocene of Timor. I. Proc. Koninkl.
Nederl. Akad. van Wetenschappen — Amsterdam. Ser. B,
74(5): 504-525.
Hummel, K. 1929. Die fossilen Weichschildkroten (Trionychia).
Eine monographisch-systematische und stammesgeschicht-
liche Studie. Geol. Pal. Abh. NF 16 (5), pp. 359-487.
Jackson, C.G. 1964. On the status of Deirochelys floridana Hay
with comments on the fossil history of the genus. Tulane Stud.
Geol. 2 (3): 103-106.
Mlynarski, M. 1969. Fossile Schildkroten. A. Ziemsen Verlag, Wit
tenberg Lutherstadt, pp. 1-128.
Nopsca, F. 1923. Kallokibotium a primitive Amphichelydian tor
toise from the uppermost Cretaceous of Hungary. Pal. Hungar.
1, 36 pp.
Parsons, T.S. and E.E. Williams, 1961. Two Jurassic turtle skulls:
a morphological study. Bull. Mus. Comp. Zool., Harvard. 125,
43-107.
Price, L.I. 1953. Os Quelonios de formacao Bauru, Cretaceo ter-
restre do Brasil meridional. Dept. Nac. de Prod. Min., Div. de
Geo. e Min., Boletim 147, pp. 7-34.
Rose, F.L. and W.G. Weaver. 1967. Two new species of Chrysemys
( = Pseudemys) from the Florida Pliocene. Tulane Stud. Geol.,
5 (1): 41-48.
Simpson, G.G. 1938. Crossochelys, Eocene homed turtle from
Patagonia. Bull. Amer. Mus. Nat. Hist. 74, pp. 221-254.
Swinton, W.E. 1939. A new fossil fresh water tortoise from Burma.
Records of the Geol. Surv. of India 74, Pt. 4, pp. 548-551.
Szalai, T. 1934. Die fossilen Schildkroten Ungams. Fol. zool. hy-
105
www.Ebook777.com
drob. 6 (2), pp. 97-192.
Watson, D.M.S. 1914. Eunotosaurus africanus Seeley and the
ancestry of the Chelonia. Proc. Zool. Soc. London. Pp. 1011-1020.
Weaver, W.G. and J.S. Robertson. 1967. A re-evaluation of fossil
turtles of the Chrysemys scripta group. Tulane Stud. Geol. 5
(2): 53-66.
__________and Rose, F.L. 1967. Systematics, evolution, and fossil
history of the genus Chrysemys. Tulane Stud. Zool., 14 (2): 63-
73.
Wegner, T. 1911. Desmemys bertelsmanni, nov. gen., nov. sp., ein
Beltrag zur Kenntnis der Thalassemydidae. Palaeontogr. Stutt
gart 58, pp. 105-132.
Wieland, G.R. 1900. The skull, pelvis and probable relationships of
the huge turtles of the genus Archelon from the Fort Pierre
Cretaceous of South Dakota. Am. J. Sci. 9, 237-251.
Wiman, C. 1930. Fossile Schildkroten aus China. Pal. Sinlca Peking
(C) 6 (2), pp. 1-56.
Williams, E.E. 1950a. Variation and selection in the cervical cen
tral articulations of living turtles. Bull. Amer. Mus. Nat. Hist.
94(9): 505-562.
__________. 1950b. Testudo cubensis and the evolution of western
hemisphere tortoises, ibid, 95 (1): 1-36.
__________. 1952. A new fossil tortoise from Mona Island, West
Indies, and a tentative arrangement of the tortoises of the
world, ibid, 99 (9): 541-560.
__________ 1953. Fossils and the distribution of chelyid turtles. 1
“Hydraspis leithii” in the Eocene of India is a pelomedusid.
Breviora, No. 13, pp. 1-8.
__________ 1954a. New or redescribed pelomedusid skulls from the
Tertiary of Africa and Asia (Testudines, Pelomedusidae). Bre
viora, No. 35, pp. 1-8.
__________. 1954b. Clemmydopsis Boda a valid lineage of Emydi-
nae Turtles from the European Tertiary. Breviora, No. 28, 9.
Wood, R.C. 1972. A fossil pelomedusid turtle from Puerto Rico.
Breviora, 392: pp. 1-12.
_____ . 1976. Stupendemys geographicus, the world’s largest turtle.
Breviora, Mus. Comp. Zool. No. 436: 1-31.
__________and M.L. Diaz de Gamero. 1971. Podocnemis venezue-
lensis, a new fossil pelomedusid (Testudines, Pleurodira) from
the Pliocene of Venezuela, and a review of the history of Podoc
nemis in South America. Breviora, 376, pp. 1-23.
Woodward, A.S. 1887. On “Leathery Turtles”, recent and fossil,
and their occurrence in British Eocene deposits. Proc. Geol.
Assoc. 10, pp. 2-14.
Yeh, Hsiang k’-uei. 1961. The first discovery of a box-turtle in
China. Vertebr. pal. Peking (1), pp. 58-64.
106
Zangerl, R. 1945. Fossil specimens of Macrochelya from Tertiary

of the Plains. Fieldiana Geol. 10 (2), pp. 1-12.
------------- 1948. The vertebrate fauna of the Selma Formation of
Alabama. Pt. 1, Introduction. Pt. 2, The Pleurodiran Turtles.
Fieldiana Geol. Mem. 3, pp. 1-56.
------------- 1953. The vertebrate fauna of the Selma Formation of
Alabama. Pt. 3, The turtles of the family Protostegidae. Pt. 4,
The turtles of the family Toxochelyidae. Fieldiana Geol. Mem.
3, pp. 61-277.
------------- 1958. Die Oligozanen Meerschildkroten von Glarus.
Schweiz. Paleont. Abh. 73, 1-56.
------------- 1969. The turtle shell. In Biology of the Reptilia, Vol. 1,
Chapter 6, pp. 311-339. Academic Press, London and New York.
------------ andR.E. Sloan. 1960. Anew specimen of Desmatochelys
lowi Williston, a primitive cheloniid sea-turtle from the Creta
ceous of South Dakota. Fieldiana Geol. 14, 7-40.
------------ and W.D. Turnbull. 1955. Procolpochelys grandaeva
(Leidy), an early Carettine sea turtle. Fieldiana Zool. 37, pp.
345-384.
107
www.Ebook777.com
Chapter 4
EMYDID TURTLES
The family Emydidae is a large and varied assemblage of

freshwater and semi-terrestrial turtles, closely related to the land
tortoises (Testudinidae) but distinguished from them by the adap
tation of the hind feet for swimming instead of walking and by the
lack of bony cores In the scales on the face of the forelimbs. The
skulls of turtles in the two families show no major differences of
structure, but emydid skulls are normally somewhat streamlined
in profile, while tortoise skulls are more upright and square-front
ed, and with a much narrower postorbital bar. Some recent authors
lump the Emydidae and the Testudinidae together into a single
huge family.
The emydid turtles, with thirty genera and some eighty-five
species, constitute by far the largest family of living turtles. The
group is almost confined to the northern hemisphere, though a few
species reach south of the equator in South America, while one
form (Pseudemys scripta dorbignyi) ventures just south of the
Tropic of Capricorn. A few species also live south of the equator in
Indonesia. The emydid turtles reach their greatest variety in eas
tern North America and in south-east Asia. There are only three
species in Europe, North America and the Middle East, though in
some places members of these species are very abundant.
The Emydidae are a modern family and have lost several shell
elements present in more primitive turtle families—for example
inframarginal and intergular scutes and mesoplastral bones. Sev
eral genera have, apparently independently, evolved a hinge in the
plastron between the pectoral and abdominal scutes, with a cor
responding breakdown in the bony structure of the bridge and the
plastral buttresses to allow flexion of the hinge to take place. How
ever, the presence of a plastral hinge is not considered taxonomic-
ally significant above the genus level, since all degrees of hinge de
velopment, from a barely-discernible flexure in the plastron which
only develops in mature turtles, to a fully flexible hinge coupled
with enlarged plastral lobes which allow airtight closure of the
shell, are found within the family. In size emydid turtles range
from the four-inch Muhlenberg’s turtle, weighing only a couple of
ounces, to the two-foot long, sixty-pound batagurs. By the way of
habitat, most species inhabit freshwater but spend many hours
108
every day basking out of the water; a few species (Malaclemys ter
rapin, Callagur bomeoensis, Pseudemys alabamensis) are usu
ally found in brackish water, while some (such as the American
box turtles of the genus Terrapene, some Rhnoclemys, Heosemys,
Geoemyda, etc.) are largely terrestrial, culminating in the ornate
box turtle, Terrapene omata, much of whose habitat is distinctly
arid. The Emydidae have recently been classified (by Samuel Mc
Dowell) into two subfamilies, based on technical features of the
rear of the skull, jaw, and cervical vertebrae. The subfamilies are:
the Emydinae (including all the New World emydids except
Rhinoclemys, and including the Old World Emys), and the Bata-
gurinae (including all the Old World emydids, plus Rhinoclemys,
but excepting Emys). The exact classification of genera, species,
and subspecies of the Emydidae is still subject to debate. The clas
sification used here is based on my own interpretation of all infor
mation available to me, but it should certainly not be considered
the last word.
109
www.Ebook777.com
A suitable starting point for a discussion of New World emy-
dids is the genus Pseudemys, which not only is the largest, most
complex, and wide-ranging of the Emydidae, but also includes the
most widely-kept pet turtle in the world, the red-eared turtle.
GENUS PSEUDEMYS
Pseudemyd turtles are medium to large in size (normal adult
lengths range from eight to about sixteen inches), and have rather
unspecialized habits, spending much time on logs or banks adja
cent to water. In diet adults are herbivorous or omnivorous. All
species have stripes on the head, neck, and limbs, though these are
sometimes lost in adult males of certain species. Males are usually
smaller than females and sometimes have several of the claws on
the forelimbs greatly elongated. These long claws are used in
courtship rituals; the male faces the female and, with hands to
gether with palms outward, caresses the female’s face with a very
rapid fluttering motion. One observer also claims that these long
claws are used for spearing worms; while this may happen on rare
occasions, it is almost certainly not a deliberate or common proce
dure.
Four basic stocks of the genus Pseudemys may be discerned;
these, named after a characteristic (or the only) species in each
group, are known as the scripta, floridana, concinna, and rubri-
ventris sections. The scripta group is characterized by the round
ed, rather than flat, ventral surface of the lower jaw (looking at
the animal from the front), by the presence of only three phalan
ges in the fifth toe, and the absence of an anterior cusp on the mid
dle ridge of the biting surface of the upper jaw. Superficially, tur
tles of •this group may be recognized by the red, orange, or yellow
horizontal line, crescent or spot behind the eye, except in certain
old males in which the pattern disappears, and the frequently
rugose and sometimes quite deeply sculptured, posteriorly serra
ted carapace. The scripta section is found over much of the south
east and south-central United States, both sides of Mexico and
Central America, the north-west comer of South America, and, in
explicably, extreme southern Brazil and northern Argentina, as
well as a small area on the north coast of Brazil. Representatives
are also found on each of the Greater Antilles, as well as on the
Cayman Islands and two of the Bahamas.
There is some debate as to which turtles of the scripta section
warrant full specific status and which should be considered only
subspecies. Clearly, where forms are allopatric but intergrade
where their ranges come in contact, we have subspecies rather
than full species. However, when taxa are isolated from each other
(such as the West Indian forms) it is probably better to accord full
species recognition. This is not an absolute rule, and elsewhere in
110
this book the isolated races of tortoises in the Galapagos are con
sidered as subspecies; it is really just a matter of one’s subjective
appraisal of the degree of difference between the forms.
The ‘typical’ race of the scripta section, Pseudemys scripta
scripta, commonly known as the yellow-bellied turtle, is found in
extreme south-east Virginia, the eastern half of North Carolina,
most of South Carolina and Georgia, and in northern Florida as far
south as Alachua and Levy counties. It inhabits a wide variety of
aquatic situations, ranging from small ponds and streams to deep,
clear springs, but is almost never seen in large basking aggrega
tions like those of Pseudemys floridana and concinna. The yellow-
bellied turtle has a short, deep, strongly sculptured, and very thick
shell; the record length is nearly one foot, but a nine-inch specimen
is considered large. Unusually large specimens are found in the
property of the Savannah River Atomic Energy project, where the
water is heated by the effluent cooling water from the atomic
reactors. The males are smaller and less domed than the females.
The yellow-bellied turtle shares most of its range with the alliga
tor, and the heavy shell may have evolved as a device to lessen the
risk of being cracked and eaten by these predators. The carapace
is basically green in the young, with a broad transverse light bar
on each of the costal scutes. The plastron is yellow, sometimes im
maculate but more usually with a black blotch on each of the gular
scutes at least, often on the humerals too, and in some specimens a
blotch is present on each of the plastral scutes. There is a broad
yellow crescent-shaped marking behind each eye, and there are
large black blotches on the underside of the marginal scutes. In
adult females the colors become less distinct, while in males they
usually disappear entirely, the dorsal surfaces of the animal be
coming black with pale areas on the anterior half that have no re
lation to the original shell pattern, while the plastron too shows in
creases in black pigmentation.
The yellow-bellied turtle lays ten to twelve eggs in May, June,
or July. In diet it is truly omnivorous; in the wild it eats aquatic
vegetation and invertebrates, while in captivity adult specimens
will eat virtually any food offered.
Occasional specimens of Pseudemys scripta scripta from the
extreme southern part of the range (in the region of Alachua
County, Florida) have highly domed shells with extremely rugose,
sculptured surfaces. It is possible that such turtles show some
genetic influence of the extinct (Pliocene) species Chrysemys
(= Pseudemys) inflata, fragments of which have been found in
Polk and Manatee Counties, further south than any member of the
scripta complex is found today. It has been suggested that the
highly rugose Pseudemys inflata bears the same relation to
modern P. scripta as Neanderthal man bears to modern man.
111
www.Ebook777.com
To the west, the yellow-bellied turtle intergrades with the Mis
sissippi Valley form, Pseudemys scripta elegans. This race,
known as the red-eared turtle, is found almost throughout Texas
and in Chaves County, New Mexico, in an as-yet indeterminate
part of north-eastern Mexico, most of Oklahoma and eastern Kan
sas, and east to Indiana, Kentucky, Tennessee and Alabama. The
area of intergradation with scripta scripta extends from Atlanta,
Georgia to Jackson County, Florida and much of southern Ala
bama. This race differs from scripta in having a lower, thinner,
and less rugose carapace, in having black blotches on all of the
plastral scutes, elongate markings on the bridge, a red streak be
hind the eye instead of a yellow crescent, and narrower transverse
bars on the costal scutes. Geographic variation exists within the
range of this subspecies; for example, specimens from southern
Texas have particularly vivid markings. The adult size is similar
to that of the yellow-bellied turtle, and the pattern of melanism in
adult males is similar. Egg complements from 5 to 23 have been
recorded, but the mean (about ten) is similar to that of the yellow-
bellied turtle, and the greater spread may simply indicate a
greater sample size. The young are carnivorous, but adults appear
to be largely herbivorous—more so than the yellow-belly. Huge
numbers of red-eared turtles are hatched in ‘farms’ in Louisiana
and Mississippi; but the capture of breeding stock for these farms,
as well as for laboratory research and demonstration, and a good
amount of just wanton destruction, do not seem yet to have made
serious inroads into the natural populations of red-ears, which re
main exceedingly common in many areas. This is a thoroughly
aquatic species preferring quiet water with a muddy bottom and a
profusion of vegetation.
In the upper portions of the Cumberland and Tennessee River
Valleys, from eastern Kentucky to north-eastern Alabama, P. s.
elegans is replaced by the Cumberland turtle, Pseudemys scripta
troosti', intergrades between the two races have recently been
found in Madison County, Kentucky. This race differs from the
red-ear in having a rather narrow yellow stripe instead of a broad
red stripe behind the eye, in having fewer but broader chin stripes,
less pigmentation on the plastron (the bridge is marked with a few
small black spots instead of with elongate dark blotches), and in
having light yellow rather than dark fingernails. The carapace is
smooth and unkeeled. The light bars on the costal scutes are some
times shortened and curled, and on rare occasions form complete
ocelli. The adult size is similar to that of elegans and scripta. No
natural history data are available for this subspecies.
In the Big Bend region of Texas, elegans gives way to the Big
Bend turtle, Pseudemys scripta gaigeae, which in Mexico extends
from northern Chihuahua southward to Coahuila and eastern
112
Pseudemys scripta gaigeae of the Big Bend Region of Texas and adjacent
Mexico. Notice the rounded spot behind the eye.
Durango. Pure individuals of this race, which are only found over
a relatively small area, are easily recognized by the large,
rounded, isolated, black-bordered orange spot on the side of the
head at the level of, but well behind, the eye. The carapace is rela
tively low and smooth and is marked with a complex reticulated
pattern which usually includes one or more ocelli on each costal
scute. The plastral pigmentation is concentrated along the mid
line, with rami extending along the seams. Near the range of ele
gans, the plastral pattern may consist of isolated black blotches.
To the east, gaigeae intergrades over a wide area with both ele
gans and cataspila.
The Sierra Madre Occidental and the deserts of New Mexico,
Arizona, Sonora, and northern Baja California constitute a hiatus
in the distribution of Pseudemys. However, a relictual population,
known as Pseudemys scripta nebulosa, survives in southern Baja
California, from San Ignacio south to San Jose del Cabo. Some
what similar to gaigeae in plastral pattern and postorbital light
spot, it differs from that race in lacking the small light spot just be
hind the eye and in having no black border to the large spot. The
carapace is short, broad, flat, keeled, and brown in ground color,
113
www.Ebook777.com
with no black ocelli, but with irregular olive reticulations. There is
a dark smudge on each marginal scute, roughly surrounded by a
squarish light mark. Males have the upwardly pointed snout
shown in several of the Mexican subspecies of Pseudemys scripta.
This is a large form; a random male and female measured by Eric
Waering (quoted by Archie Carr) measured 13.0 inches and 14.6
inches in straight length, respectively. There is a north-south cline
in certain characters, southern representatives having black in
stead of brown plastral markings, olive instead of brown ground
color, and an orange temporal spot instead of a yellow one. This
variation is not surprising considering the isolation that must exist
between each deme of this race; the habitat is mainly disconnect
ed pools and lagoons separated by large areas of impassable
desert.
Another very large form, P.s. hiltoni, is found across the Gulf
of California in the Lower Rio Fuerte of southern Sonora and nor
thern Sinaloa. It is distinguished from nebulosa by the presence of
intensely black, smudge-like spots on the upper and lower surfaces
of each of the lateral and posterior marginal scutes. The plastral
figure shows some similarities to that of gaigeae, but consists of a
large black blotch covering almost all of the median halves of each
of the plastral scutes except the gulars. The bridge is also marked
by a solid black blotch, which may show a break at the pectoral-
abdominal seam. The supratemporal stripe is light orange in
color.
To the north, in the Sonora, Yaqui, and Mayo drainages of
coastal Sonora, hiltoni is replaced by the newly-described Pseu
demys scripta yaquia. In this race the plastral pattern almost dis
appears in adults, but the vestiges which may be apparent are in
the form of narrow lines clustered around the midline and the
seams of the plastron, rather than large black blotches. The post
orbital mark is yellowish-orange; the lateral scutes have jagged
black blotches at the centers of poorly defined ocelli. P.s. yaquia
inhabits a wide variety of permanent and temporary aquatic situa
tions, and Legler and Webb, describers of the race, consider that
man’s hydrological works in northern Mexico may have created
more consistently favorable habitat than formerly existed.
Many authors have written that the race Pseudemys scripta
omata extends all the way from Mexico (both drainages) to South
America. It now appears, however, that ornata in the strict sense
is restricted to an area of western lowland Mexico, extending from
the Rio San Lorenzo drainage of Sinaloa to somewhere between
Tuxpan, Nayarit and Acapulco, Guerrero; the type locality is Maz-
atlan. I have not seen Pseudemys from this area, and we must
await Legler’s redescription for details of it. However, from the
key to north-western subspecies of P. scripta provided by Legler
114
and Webb in their description of Pseudemys scripta yaquia, it ap

pears that true omata is characterized by a carapace of a dark
brown ground color, each costal scute bearing a distinct ocellus
consisting of 2 or 3 pale concentric lines and a darker center. The
ocellar pattern is never lost in adults. The dark centers of the ocelli
do not contrast sharply with the ground color. The plastral pattern
is distinct, consisting of four concentric, dusky lines on a pale yel
low ground, fading somewhat with age but never completely ob
scured in adults.
Further south still, the race Pseudemys scripta grayi is known
from the Pacific slopes of Oaxaca, Chiapas, Guatemala, El Salva
dor, and probably Honduras. As with all the Pacific Pseudemys,
this race reaches a large size; a 14.75” carapace I was given in
Tegucigalpa, Honduras, was not considered uncommonly large by
the local people. P.s. grayi (also frequently known as P.s. umbra)
has a rather plain, almost uniformly brown carapace with a single
more or less circular black blotch near the posterodorsal comer of
each costal scute, and at the junction of each pair of marginal
scutes. These black blotches are sometimes enclosed within very
poorly defined light ocellus. The carapace is moderately rugose,
Pseudemys scripta grayi of the Pacific drainage of southern Mexico to El

Salvador.
115
www.Ebook777.com
very thick, and in fully adult individuals has a somewhat oblong,
straight-sided shape. The ventral markings include dark-centered
submarginal blotches, a dark linear pattern on each bridge, and
areas of dark pigment concentrated toward the anterior ends of
each of the plastral scutes except the gulars. The extremities are
of a dark grey-brown color, the head and neck being adorned with
numerous rather narrow yellow lines but without any expanded or
erythristic temporal bar. As with all Pacific Pseudemys, the jaw
edges are markedly serrate. The nostrils of mature males are per
forated distinctly below the tip of the snout. In very young speci
mens, the yellow head lines are relatively much broader and the
plastron bears a pattern of concentric curved lines concentrated
along the midline. This turtle is highly appreciated for meat
throughout its range, and turtles of this subspecies were the most
expensive I saw offered for sale in Guatemala City. In Chiapas it is
reported by Miguel Alvarez del Toro to live in lakes and sluggish
rivers and to lay clutches ranging from ten to twenty eggs from
February to April. Del Toro reports this turtle to be particularly
abundant in the vicinity of Acapetagua, Chiapas. It is uncertain
whether grayi intergrades with venusta across the Isthmus of
Tehuantepec. Moll and Legler, in their range map for the species,
show no contact between the Pseudemys of the Gulf drainages and
the Pacific drainages in this region. On the other hand, Gustavo
Casas Andreu, in his thesis ‘Las tortugas dulciaguicolas de
Mexico,’ shows a range map with the species reaching clear
across the isthmus.
A race of Pseudemys scripta is found in Lake Managua and
Lake Nicaragua and the rivers draining into these lakes in Nicara
gua; however, this population has yet to be properly described,
and no subspecific name is currently available for it. Yet another
undescribed race is found from the Rio Terraba drainage of Paci
fic Costa Rica to the Rio Bayano and Chucunaque drainages of
eastern Panama.
Returning to the Gulf Coast of Mexico, P.s. elegans is replaced
in coastal Tamaulipas by P.s. cataspila, whose range extends to
the south to the vicinity of Punta del Morro, Veracruz. The exact
northern border of the range is unknown; northern Tamaulipas is
so arid that the range is probably highly discontinuous in this area,
but it is known that inland pure elegans is found as far south as
Ciudad Victoria. P.s. cataspila is considerably larger than ele
gans ; a female I found near Rancho Nuevo, Tamaulipas, has a 13”
carapace. The carapace is slightly rugose and moderately domed,
with a highly streamlined profile. Young specimens are vividly
marked with an orange ocellus enclosing a large black blotch typi
cally located in the posteroventral corner of each costal scute.
However, in many specimens the ocelli tend to be incomplete and
116
Specimen of Pseudemys scripta elegans from Ciudad Victoria, Mexico, the

southernmost limit of the known range.
to have branching tails, and to be located near the middle of the

posterior border of the costal scutes. Each marginal scute is deco
rated with a partial ocellus which encloses a black spot. The cara
pace is otherwise some shade of green, with various ill-defined
light reticulations.
The plastron in young cataspila has the central pattern of con
centric seam-following black lines typical of many races of tropi
cal scripta. However, this pattern fades as the turtle grows and is
replaced by blotchy stretches of intensely black pigment along the
plastral seams, which may form large areas of unrelieved black
towards the rear of the plastron. The linear pattern on the bridge
and the submarginal blotches similarly become intensified and
smudgy. This secondary pigmentation, however, fades out as the
turtle matures, and big old females have virtually no dark pigmen
tation anywhere on the ventral aspect of the shell.
The head pattern of cataspila includes a moderately expanded
temporal bar and numerous longitudinal yellow lines; two broad
yellow lines extend downward and backward from each nostril,
and another, broader still and often of a slightly orange color, ex
tends downward and backward from the rear corner of the orbit.
Poorly contrasting olive lines extend longitudinally along the
limbs.
117
www.Ebook777.com
Mature cataspila may show melanism in the carapace. This
commences with a blackening of the scute borders and may ulti
mately result in extensive breakup and discoloration of the dorsal
pattern, though some trace of the costal ocelli usually remains.
The habits of this race have not been studied.
There are two internally drained basins in northern Mexico
which have populations of Pseudemys scripta. One of these is the
Rio Nazas drainage in Coahuila and Durango, and the other is the
basin of Cuatro Cienegas in central Coahuila. The first population
has yet to be described; J.M. Legler and E.O. Moll have intimated
that it is distinct, though related to P.s. gaigeae. The second has
been called Pseudemys scripta taylori. Seven points were cited in
which taylori differed from elegans; however the comparison with
the form from the coast of Tamaulipas, here called cataspila, was
based on only four specimens, and in fact this coastal race was
summarily, though ‘tentatively’, dismissed as indistinguishable
from omata (in the old, inclusive sense). In fact taylori is probably
a synonym of cataspila; certainly none of the characters cited by
Legler serve to distinguish these two races.
South of the City of Veracruz, cataspila is replaced by Pseu
demys scripta venusta, whose range extends along the humid
coastal lowlands of the Gulf Coast through the States of Tabasco
and Campeche, and across the moist southern part of the Yucatan
Peninsula to Belize. In this race the carapace is adorned with large
dull orange ocelli on each of the costal scutes. These ocelli are
located toward the posterodorsal comer of the scutes, and each en
closes a black smudge. Linear or curved markings, also dull
orange in color, are present on the central scutes, and the margi
nals have partial ocelli enclosing black spots. However, the overall
shell pattern is much simpler than that of cataspila, with far less
vermiculation. Ventrally, submarginal blotches are present at the
junction of each of the marginal scutes and there is an elongate lin
ear figure on each bridge. The plastral figure consists of a more or
less symmetrical pattern of dark lines concentrated along the mid
line and reaching laterally wherever it crosses a seam. The extent
of this plastral figure is extremely variable. The pattern may fade
with age, but melanism does not normally occur in this subspecies.
The head pattern consists of several strong yellow lines extending
posteroventrally from the nostrils and eyes; on the other hand, the
upper surface of the head has only the faintest linear markings. A
broad temporal bar is present but is so dark that it barely shows
against the dark ground color of the head. The front face of each
forelimb is marked with one broad and one narrow yellow stripe.
The rump is marked with a pair of horizontal lines which extend
from the hind feet to the tail. The tip of the snout of mature males
is strikingly elevated, with the nostrils perforated below the tip.
118
Females have broader heads than males and lack the terminal ele
vation of the snout.
I am not certain of the maximum size reached by this race.
The numerous specimens seen in the markets of Veracruz are
rarely more than about eleven inches in length. On the other hand,
Alvarez del Toro reports that in Chiapas this race can reach a
length of almost half a meter and a weight of about ten kilograms.
Del Toro also reports that in Chiapas nesting takes place at night
from January to April, and on certain river beaches nests may be
made in such density that hundreds of eggs can be gathered in a
few minutes. The eggs are reported to number between twelve and
twenty per clutch and to measure approximately 1.6” x 1”. The
nest cavity is about eight inches deep. This turtle is of considerable
commercial importance and is eaten in large numbers throughout
its range.
Turtles of the species Pseudemys scripta are found in isolated
waterholes in the arid northern part of the Yucatan Peninsula.
However, this population has not been described, nor its racial
characters determined.
From northern Honduras along the Caribbean drainages of
Nicaragua, Costa Rica, and Panama, and in the Rio Atrato drain
age of northern Colombia, venusta is replaced by a closely related
subspecies, still lacking a name, distinguished by its somewhat
light coloring, the presence of a row of small ocelli on the rump,
and the presence of elongate loops rather than continuous lines
under the neck. The head shows the same sexual dimorphism as
venusta. The temporal bar is narrow but clearly defined, and is
yellow or orange in color. This subspecies has recently been
studied in detail in Panama by Moll and Legler (1971). These
authors concluded from their findings that Pseudemys scripta is a
relatively recent immigrant to the tropics and shows only the mini
mal adjustment to a tropical life. Thus, nesting is still highly sea
sonal (December to May, i.e. the dry season), and relatively large
clutches are deposited (nine to 25 eggs, one to six clutches per sea
son). This reproductive behavior contrasts with that of the sym-
patric Rhinoclemys and Kinostemon species, which tend to pro
duce very small clutches almost throughout the year. As with
other races of P. scripta, the nests of the Panamanian subspecies
are subject to such intensive destruction, especially by birds,
Ameiva lizards, and armadillos, that it is amazing that the turtles
manage to survive at all, let alone to maintain their high popula
tion numbers. P. scripta in Panama is largely vegetarian, but
about half of the stomachs examined contained at least some
animal matter.
A further subspecies, P.s. callirostris, is locally common in
extreme northern Colombia and north-western Venezuela. This
119
www.Ebook777.com
race has ocelli on the costal scutes, though these may be interrup
ted by the hind margin of the scutes, and an extensive and very
decorative plastral figure consisting of broad greenish or black
lines, sometimes continuous with the lines on the bridge, and en
closing various circles, ovals, and irregular curved figures. The
race is immediately recognizable by the presence of enclosed, cir
cular light markings on the chin and jaws. There is a broad, pale
red postorbital bar which does not enter the eye. This subspecies
appears not to reach a large size; I have never seen one more than
about ten inches long. Only a few years ago large numbers of
hatchlings of these attractive turtles were being shipped to the
United States for the pet trade.
A gap of thousands of miles separates callirostris from the
terminal member of the scripta series, Pseudemys scripta dorbig-
nyi, a form which is frequently accorded full specific status (Pseu
demys dorbignyi). The presence of this population in Rio Grande
do Sul, Brazil and adjacent parts of eastern Uruguay and northern
Argentina (Entre Rios Province) is totally unexpected and hard to
explain zoogeographically. Marcos Freiberg has recently
proposed that the Brazilian population, in the Rio Guaiba, is
worthy of subspecific recognition (Pseudemys dorbignyi brasilien-
sis). This species has been reported recently from northern Brazil,
near Sao Luis, Maranhao, though whether or not this population is
Pseudemys scripta dorbignyi from Argentina. If recognized as a full species,

this would be P. dorbignyi dorbignyi.
120
introduced should be investigated, dorbignyi being sold in the pet

trade in Brazil in the same way that elegans was in the United
States. Pseudemys scripta dorbignyi has a plastral pattern similar
to that of callirostris, and many of the head markings are also
similar, but the chin spots are reduced to a single elongate mark
ing beside each comer of the lower jaw. Males may become exten
sively melanistic, losing all traces of the juvenile pattern.
The races may be distinguished by several characters. The
adult female of dorbignyi has a general dark brownish hue with
very intense reddish-orange markings, the supratemporal stripe is
not very broad, and the plastron has a large central black marking
on a reddish orange background. The female of brasiliensis, on the
other hand, has an olive-green aspect with yellow or slightly rosy
markings. The supratemporal stripe is very wide, and the black
plastral markings have a yellow background. The males are dis
tinguished by the unmarked light olive-green plastron of brasilien
sis, which contrasts with the black and yellow plastron of
dorbignyi. There are also some reported differences in scutellation
and plastral dimensions; dorbignyi has an almost square first
vertebral scute, approximately equally wide anterior and pos
terior plastral lobes, a marked overlap of the outer border of the
femoral scutes over the anal scutes, and an interfemoral seam
that is usually less than the length of the interanal seam. In brasili
ensis the hind margin of the first vertebral is notched, the
posterior lobe of the plastron is wider than the anterior lobe, there
is minimal overlap of the femoral scute border over the anal
scutes, and the interfemoral seam is equal to or greater than the
interanal seam. The juveniles are said to be distinguished by the
more brilliant coloration of brasiliensis and the presence in the
latter of relatively simple angular or three-pointed-star markings
on the laterals; juvenile dorbignyi have semicircular markings on
the laterals, with the open side directed backward.
Most if not all of the characters for distinguishing the two sub
species are of the type that tend to be very variable, and later work
may show at least some of them to be unreliable.
Rather few data are available on the life history of these
southern cooters; we are indebted to Marcos Freiberg for most of
the available information. According to Freiberg, brasiliensis
reaches a carapace length of about 26 cm and feeds upon small
fish; in captivity, the species is carnivorous, consuming pieces of
beef avidly. They hibernate in winter and during the warm months
may form large basking aggregations. One female laid fourteen
eggs during a fifteen-day period in June; the eggs measured 41x27
mm and had the usual tough, flexible shells.
In the West Indies, Pseudemys is present (and is the only
freshwater turtle) in Cuba, Jamaica, Grand Cayman, Cayman
121
www.Ebook777.com
Brae, Hispaniola, Puerto Rico, Vieques Island, and Great Inagua
and Cat Island, in the Bahamas. Some of these island races show
great similarity to Pseudemys scripta grayi and almost certainly
reached the islands from Central America rather than from Flori
da, where Pseudemys scripta even today barely enters the penin
sula. Most of the island races show two phenomena typical of
many island animals: considerable individual variation and a loss
of distinctive color patterns. The former is the result of a lack of
competition; a freshwater turtle niche exists in the islands, and
there are no other species of turtle competing for it. Consequently
rigid specialization is unnecessary, and no ‘ideal type’ exists
which is better adapted than a slightly different type. The color
pattern is liable to break down in island populations because spe
cies distinguishing marks are unnecessary where no other turtle
species are present; any female turtle that a male turtle can find is
suitable for mating. On the other hand, in areas where a number of
turtles species occur in the same habitat, it is vitally important
that females of each species carry distinguishing marks by which
males can recognize their own subspecies. A corollary of this is
that turtles, such as the West Indian Pseudemys, which do not
share their habitat with other turtle species will mate with turtles
of completely unrelated types in captivity, while turtles which are
found in nature coexisting with many other species will usually as
siduously seek out females of their own species when kept in cap
tivity in a large mixed collection.
A uniform characteristic of the island turtles, shared by some

tropical mainland members of the genus but contrasting sharply
with most Pseudemys races in the United States, is their aggres
sive disposition and tendency to bite when picked up. This snap-
ping-turtle-like nature may indicate that the island forms are
more active predators that the largely vegetarian North American
races.
Pseudemys decussata, the jicotea of the Cubans, contains

three subspecies: decussata from most of Cuba and the Isle of
Pines; plana from the Rio Jobabo, in western Oriente, Cuba; and
angusta, from the Taco River, Pinar del Rio, Cuba. The typical
form has a rather broad head and moderately domed carapace,
with a keel present at the rear. The skin is greenish, with white or
yellowish stripes; a pinkish line extends from behind the eye. The
melanistic male is of an overall greenish-yellow color, with very
strong, extensive black vermiculations on the carapace and plas
tron and, to a lesser extent, on the limbs. This form is found mainly
in sluggish rivers with deep pools, though it does not scorn other
types of bodies of fresh water. It is carnivorous, eating molluscs,
crayfish, etc. It is said to be very wary and difficult to catch.
122
The Cuban Pseudemys decussata. Photo by Dr. W. Sachsse.
The plana variety is rather similar, but the carapace is broad

and very flattened. The plastron is similar, that is, dusky, with a
curved symmetrical linear figure, well-defined only in the juve
nile. The postorbital stripe is bright red, but the other lines, below
and parallel to the postorbital, are yellow. The carapace is brown
and somewhat reticulate. The race angusta has a narrower and
more elongate carapace than the typical form; the sides of the
carapace are straight, or even slightly constricted. The ground
color of the soft parts is darker, and the limb stripes are more dis
tinct. The upper jaw is strongly notched; the top of the head is dark
brown. The plastron is light yellow, with portions of a dusky figure
usually present, following the seams anteriorly, but extending
over the posterior scutes. The melanistic male has the hind half of
the carapace solid black and the soft parts blacker than in a mela
nistic example of P.d. decussata. A specimen of this turtle with a
carapace slightly more than 15 inches in length, and weighing 22.5
pounds, has been found.
Pseudemys terrapen, found all over the island of Jamaica ex
cept for the north coast, has a long, rather low wedge-shaped cara-
123
www.Ebook777.com
pace of a brown or brownish-grey color. There may be faint dark
blotches on the gulars and humerals. The color pattern of the soft
parts is almost lost, but faint greenish-yellow lines are present on
the neck and limbs. The snout is slightly conical and projects be
yond the jaws. The juvenile has an extensive dark, solid pattern on
the plastron, covering all except an Indented marginal area. The
carapace length may exceed one foot. This species differs from the
Cuban forms in having denticulated rear marginals, and from the
Cayman Islands form by the more flaring hind carapace margin.
Certain large male specimens from riverine situations in southern
Jamaica are reported to assume an overall white color. A female
dissected on April 14 contained four shelled eggs, each about 1” x
1.5” in size, and four large yolks. Males have claws about four
times as long as those of the female, and these are used in the typi
cal courtship rituals; however, instead of tickling the face of the
female as is usual, the male actually places the forefeet, palms
outward, on each side of the female’s face and precedes with a
rapid slapping movement, slapping harder on the side to which the
uninterested female attempts to turn.
Pseudemys felis, found only in the vicinity of Tea Bay, Cat
Island, Bahamas, is very similar to P. terrapen and may conceiv
ably be an introduced population of that species; certainly, if the
colonization of Cat Island was natural, it is strange that no Pseu
demys populations became established on any of the intervening
islands. However, felis is supposed to have a shorter, higher, more
circular carapace than terrapen, with a slight keel. The limbs and
head are grey, and the skin pattern is, if anything, even further
gone than in terrapen. This species lives in temporary pools near
Tea Bay; when the pools dry up in the hot season, the turtles bury
themselves. They are said to be very fond of a local fruit known as
the custard-apple, which the natives use as bait to catch these tur
tles. This species exhibits a certain degree of melanism, though
mainly in old males.
The only other Bahamian Pseudemys is P. malonei, which is
locally abundant in the very few freshwater ponds on Great Inagua
Island. Young males have a purplish-brown, unmarked carapace
and a plastron of a dirty yellow color with looping black lines along
the seams. This race is somewhat similar to the nearby form on
Hispaniola, but the soft parts differ in having a unique purplish
tinge. All the normal head stripes of mainland Pseudemys are pre
sent, but are poorly contrasting and are only visible under bright
light. The shell is quite low in young specimens but may be very
high in big females. Old males become strongly melanistic, as
suming the so-called ‘rugosa’ condition (Pseudemys rugosa is an
invalid name based on a melanistic example of one of the Antillean
terrapins).
124
Two forms have been described from Hispaniola: Pseudemys

decorata and Pseudemys stejnegeri vicina. However, from look
ing at several dozen Pseudemys from Haiti I am inclined to the
view that the two names refer to a single form. Typical vicina
have a yellow plastron with a pattern of curved lines, often form
ing irregular ocelli along the seams. The carapace may be pale
yellow-green to almost black, while in melanistic individuals the
pattern of uniform black on the posterior half of the shell, and
black mainly confined to the seams on the anterior half, may be
expressed. Decorata does not appear to get melanistic, the cara-
Variation in plastral markings of the Hispaniolan Pseudemys decorata
125
www.Ebook777.com
pace being usually pale olive-green. The plastron is marked with
small black spots, asymmetrically placed. The postorbital bar is
rather narrow, and is usually yellowish or greenish, but occasion
ally red. The forms are reportedly sympatric, and a series of indi
viduals shows all intermediates between the ‘decorata’ pattern
(which has no duplicate on any other island), and the more usual
‘vicina’ pattern. Moreover, the differences in plastral pattern ap
pear to be associated with sex, and a separation of a series of spe
cimens into ‘vicina’ and ’decorata’ patterns also tends to separate
the males from the females. The Haitian turtle reaches a length of
about one foot. The single specimen I have seen from the Domini
can Republic (a male) had spots on the plastron, an acutely point
ed snout, and a somewhat deeper, narrower carapace than is nor
mal for Haitian specimens.
The Puerto Rico form, Pseudemys stejnegeri, barely exceeds
nine inches in maximum carapace length. It is recognized by the
unmarked reddish-brown carapace and the constriction at the an
terior ends of the humero-gular seams. A pattern is present on the
plastron, being distinct anteriorly and extending on to the upper
surface of the gulars, but becoming less distinct posteriorly. The
skin is brownish with yellow stripes. The postorbital stripe is usu
ally bright red. This race is also found on Vieques Island, near
Puerto Rico.
Pseudemys granti is found all over Grand Cayman Island and
is introduced but uncommon on Cayman Brae; it is locally known
as ‘hick-a-tee,’ an anglicization of the Spanish ‘jicotea.’ The very
young have ocelli on the costal scutes, a pale orange postorbital
bar, and a plastral pattern somewhat intermediate between the
linear and the spotted type; the general effect is of a rather
bleached-out grayi. Adult females reach a length of about a foot
(males about eleven inches) and have stout, moderately flattened
and longitudinally rugose shells. The nuchal scute is distinctly nar
rower than that of terrapen. The male is slightly flatter than the fe
male, has long claws on the forelimbs, and in the usual melanistic
form has a light grey shell with black lines along the seams and
small black spots on the carapace. The female gradually loses the
juvenile pattern with maturity, the shell becoming uniformly
dark, but traces of the head striping remain.
The other three sections of the genus Pseudemys are virtually
confined to the United States. The floridana group, of the south
eastern States, is identified by the immaculate plastron, domed
shell, and the relatively smooth rear margin of the carapace (ser
rated in scripta). These are large turtles and have the highest
point of the carapace anterior to the mid-point. The bridge is very
deep in lateral profile. They lack the thick epiplastral lip shown by
the United States races of scripta, and neither sex becomes mela-
126
nistic. Adults appear to be entirely vegetarian. The jaws are

slightly serrated, but (except in the Missouri slider) there is no
prominent notch at the front of the upper jaw, as there is in the
rubriventris group and some of the western races of concinna.
The Coastal Plain turtle, Pseudemys floridana floridana, is
found along the coastal strip from extreme south-eastern Virginia
to south-east Alabama and northern Florida. This turtle is found in
rivers, swamps, and ponds; it reaches a maximum length of 15.6
inches. The plastron is yellow with no greenish tint; the bridge is
marked with a series of light-centered dark blotches, situated on
the seams between the marginals. The ground color of the soft
parts is brown, and the stripes yellow. The two stripes along the
sides of the top of the head (the paramedian and supratemporal
stripes) do not join. This is the commonest turtle of the Okefenokee
Swamp in south-eastern Georgia. Eggs are laid in May and June;
clutch sizes from twelve to twenty have been recorded. The eggs,
like those of all Pseudemys, are elongate and slightly soft-shelled.
In peninsular Florida, the Coastal Plain turtle is replaced by
the peninsula turtle, Pseudemys floridana peninsularis. This is one
of the commonest and most conspicuous turtles in Florida, and is
found throughout the peninsula except for the Everglades area.
This race is distinguished from floridana by the greenish tint of the
plastron and striping of the soft parts. Also, the ground color of the
head is intense, sooty black, and the paramedian and supratem
poral stripes join above and behind the eyes, forming a character
istic ‘hairpin’ configuration. The largest specimen on record, from
Orange County, Florida, had a shell length of 15.8 inches. Such
huge females develop a distinct shoulder in the shell above the
forelegs and frequently also a highly domed carapace with a dis
tinct median depression. Nesting has been recorded in all months
from November to June, and it is probable that an individual turtle
nests at least twice during one year. Recorded clutch sizes range
from 12 to 29 eggs; frequently one or two of the eggs are laid a few
inches away from the main clutch in shallow side pockets. A tre
mendous proportion of nests is destroyed by skunks, raccoons,
possums, king snakes, hogs, and, in recent years, armadillos.
To the west the Coastal Plain turtle is replaced by the Missouri
slider, Pseudemys floridana hoyi; this race ranges from southern
Alabama north to extreme southern Illinois, and west to eastern
Kansas, Oklahoma, and Texas. This race is smaller than its eas
tern counterparts, the record length being 11.8 inches. The plas
tron is immaculate yellow or orange. There Is a notch at the front
of the upper jaw, but this is not flanked by sharp cusps, and the
jaws are not serrate. The shell is relatively short and broad and
often somewhat oblong in shape. The carapace is brownish or olive
with numerous light, concentrically arranged curved lines. The
127
www.Ebook777.com
Turtles of the species
Pseudemys floridana
have no plastral mark
ings.
head is decorated with a large number of narrow lines, mostly

longitudinally arranged, approximately equal in width, and fre
quently broken and asymmetrical, at least dorsally. There are in
dications that this race is more carnivorous than its eastern rela
tives, though Strecker’s remarks that in nature it feeds almost en
tirely on molluscs is oddly at variance with the thin, delicate jaws.
The concinna section of Pseudemys is distinguished from flori
dana by the dark markings along the plastral seams and the usual
ly lower, more streamlined shell profile. The river cooter, Pseu
demys concinna concinna, is found from southeast Virginia
through almost all of North and South Carolina and Georgia to the
eastern edge of Alabama, where intergradation with the hierogly
phic turtle takes place. The river cooter hybridizes with the Coas
tal Plain cooter in many areas. This is a relatively small form, the
record shell length being 12.75 inches. The shell is long and depres
sed and the carapace markings include a light ‘C’ on the second
costal scute, amid a welter of complex concentric patterns and
128
angular ocelli. The plastron is yellow, sometimes with a reddish

tinge, and there is a dark figure more or less confined to the
seams, at least anteriorly. There are five to seven parallel lines on
the top of the head between the broad supratemporal stripes. The
eyes are a most attractive turquoise color. Little information on
the diet of this race is available, though filamentous algae have
been found in stomachs. It usually lives in rather fast-running
streams and rivers.
To the west, the river cooter intergrades with the hieroglyphic
turtle, Pseudemys concinna hieroglyphica, whose range extends
north to extreme southern Illinois, and west to eastern Texas,
Oklahoma, and Kansas. This is a flat, streamlined turtle, and the
rather narrow shell is often constricted at the region of the sixth
marginals. The carapacial figures are composed of more, and nar
rower, lines than are those of concinna, and are rounded rather
than angular. Also the size is larger, the record length being 14.6
inches. This race shows a preference for quiet backwaters of
rivers, and is rarely found in fast-moving water.
Head of
Pseudemys con
cinna hieroglyph
ics.
Along the Gulf Coast from the panhandle of Florida to Louisi

ana and Texas, the hieroglyphic turtle intergrades with the Mobile
turtle, Pseudemys concinna mobiliensis. This race has a higher
shell than concinna and hieroglyphica, and is very similar to flori-
dana except that it has plastral markings—the usual pattern of
dark lines along the seams. The maximum recorded shell length is
13.75 inches, but there is evidence that larger individuals exist.
129
www.Ebook777.com
This turtle is found in coastal streams, preferably those with
abundant submerged vegetation, in brackish estuaries, and occa
sionally on saltwater flats behind the offshore islands. On the Gulf
Coast of Florida, from Franklin County to Pinellas County, the
Mobile turtle is replaced by the Suwannee turtle, Pseudemys con-
cinna suwanniensis. This is the largest member of the genus in the
United States, reaching a record length of 16.3 inches. This race is
darker in color than mobiliensis, many specimens appearing
almost black; moreover the stripes on the skin are greenish-white
rather than orange-yellow, and the number of stripes on the head,
neck, and limbs is reduced. The plastron is yellow or orange, and
except in old, faded specimens shows the usual pattern of dark
markings along the seams. This turtle is abundant in the Suwan
nee and Santa Fe River Systems, and in a number of the clear
Florida springs it is found in association with peninsularis, though
hybridization is very rare. It also ventures out into the Gulf of
Mexico from the mouth of the Suwannee River. This race appears
to be completely vegetarian, at least when mature. In parts of its
range it reaches great concentrations, but the huge basking aggre
gations seen even fifteen years ago are no longer seen today, great
numbers of these turtles having been caught for their delicately-
flavored flesh.
The Suwannee turtle apparently has completely cryptic breed
ing habits, which is all the more amazing since other Florida Pseu
demys are frequently found nesting. Indeed Crawford Jackson,
who conducted an intensive investigation of this turtle for his
Ph.D. dissertation, was obliged to remove eggs from the oviducts
of a female turtle and incubate them in order to obtain hatchlings.
It is also strange that Suwannee turtles are literally never encoun
tered crossing highways, although practically all other Florida
turtles (except Stemotherus and Macroclemys) are found on high
ways more or less regularly during suitable weather conditions.
The only remaining member of the concinna complex is the
Texas slider, Pseudemys concinna texana, which is found in most
of southern Texas, as well as the northern parts of Chihuahua,
Coahuila, Nuevo Leon, and Tamaulipas; in eastern Texas it inter
grades with hieroglyphica and mobiliensis. This is a most distinc
tive race. The plastral pattern in the juvenile consists of narrow,
clearly-defined lines along the seams; these are lost in the adult.
The head pattern is exceedingly complex and variable and con
sists of numerous broken lines. The upper jaw is cusped—a unique
feature in this species. The carapace of the juvenile is beautifully
marked with a complex pattern of loops and whorls; in the adult
this pattern is lost. This race is not known to exceed a carapace
length of 10.75 inches. It is most commonly found in rivers, but is
sometimes encountered in ditches and cattle tanks.
130
Plastron of
Pseudemys con
cinna texana from
southern Texas
and northeastern
Mexico.
Turtles of the rubriventris section of Pseudemys are identified

by the notched, cusped upper jaw and the pink or reddish tinge to
the plastron. The carapace markings too are red in half-grown and
adult specimens, but are yellow in hatchlings. Four isolated popu
lations exist, which are referable to three species. Pseudemys
rubriventris, the red-bellied turtle, is found in a rather confined
area extending from the southern half of New Jersey inland to the
eastern tip of West Virginia and south to extreme eastern North
Carolina. A completely isolated colony exists in Plymouth County,
Massachusetts, but although this population was formerly consid
ered to constitute a separate subspecies (bangsi), Conant has now
shown that the supposed differences do not hold up when adequate
series are used for comparison. The Florida red-bellied turtle,
Pseudemys nelsoni, is found throughout the peninsula of Florida,
while the Alabama red-belly, Pseudemys alabamensis, is found in
the vicinity of Mobile Bay, Alabama and possibly also Apalachee
Bay, Florida.
Pseudemys rubriventris has a long, rather low carapace with
the highest point near the middle. Large females are about ten or
eleven inches long, while the record length may be over fifteen
131
www.Ebook777.com
A very dark red-
bellied turtle, Pseu-
demys rubriventris.
inches. The carapace is somewhat rugose and is brownish in

ground color, with red transverse bars and many other smaller
red markings. The plastron is orange or red, with a complex figure
composed of elongate dark blotches in the young, but without such
a figure in adults. Old individuals become very dark, but this mel
anism, unlike that of certain Pseudemys scripta races, does not
obliterate the head markings. This turtle is found in ponds, lakes,
and streams and, although very fond of basking, is very shy and
hard to catch. The laying season varies with latitude, being early
June to July in New Jersey but from late March to early June in
Virginia. One old writer mentioned that as many as thirty-five
eggs may be laid, but this has not been corroborated since, and in
dividuals big enough to lay so many eggs are very hard to find
nowadays.
The Florida red-bellied turtle, Pseudemys nelsoni, is abundant
throughout the peninsula of Florida and is especially common in
the Everglades (where the peninsula turtle is absent). It has a
deeper and much heavier shell than rubriventris, and the ground
color of the soft parts is intense black. The markings of the lower
marginals are solid (ring-shaped in rubriventris), and the hatch
lings have only a few small black spots along the midline of the
132
plastron, which are rapidly lost with growth. The bridge itself is
usually unmarked, while that of rubriventris bears a heavy black
band. The head markings also are fewer than those of rubriven
tris, but the median stripe along the snout and the crown of the
head is usually very prominent and has backwardly-directed rami
extending from the anterior point, producing a prefrontal 'arrow.’
The carapace of adult specimens is black, with a broad red trans
verse bar across each of the first three costals. It is possible that
this turtle has gained in relative abundance during the last twenty
years; Carr found it to be much rarer than peninsularis in all habi
tats where they were found together, yet today it appears to be just
about as plentiful as peninsularis. It is found in the clear Florida
springs, as well as in ditches, lakes, sloughs, and marshes. It is
probable that the nesting season extends throughout the year. In
diet the Florida red-belly is predominantly herbivorous, showing
no real differences in dietary preference from the peninsula turtle,
with which it shares most of its range.
The Alabama red-bellied turtle, Pseudemys alabamensis, is
still known from so few specimens that an accurate diagnosis of
the species cannot be given; indeed, for many years, the species
was considered invalid, it being argued that the type specimen was
simply a Mobile turtle (Pseudemys concinna mobiliensis) with an
aberrantly cusped jaw.
The Alabama red-bellied turtle is known with certainty only
from the vicinity of Mobile Bay, Alabama, where it is found both in
rivers and in the salt waters of the bay itself. However, there are
reports of specimens both from Apalachee Bay, Florida, and from
eastern Texas, and if these prove valid the species may have a
fairly wide, though almost certainly discontinuous, range along
the northern shore of the Gulf of Mexico. The species is obviously
closely related to P. nelsoni, from which it differs in having less re
duction of the light striping on the head. The strongest head stripes
are: i) a broad horizontal line along each side of the neck, which
forks below the corner of the mouth, sending one ramus up toward
the eye and the other along the lower jaw; ii) a broad median band
which extends back from the symphysis of the lower jaw, branch
ing posteriorly into two rami which run parallel along the under
side of the neck ; iii) a stripe of moderate width which extends pos
teriorly and ventrally from the upper rear comer of the eye; iv)
converging supratemporal stripes that run along the temporalis
muscles as far forward as the orbits; v) a strong median dorsal
stripe between the eyes that tapers anteriorly to form a light pre
frontal ‘arrow.’ In addition, thin stripes run between and parallel
to each of the bold stripes listed above.
The carapace of P. alabamensis is rugose ; it is highly arched,
reaching its highest point anterior to the middle, and its widest
133
www.Ebook777.com
point at the middle. The nuchal scute presents a long dorsal
aspect, but only a short ventral aspect or ‘underlap.’ The carapace
is olive to black in younger specimens, with red to yellow bars on
the costals and marginals; however, older specimens of both sexes
show a breakup of the dorsal pattern into a complex vermiculation
of red-brown and black. The plastron is usually reddish, but it too
may become vermiculated with black in adults. The skin is olive to
black, with yellow stripes. The lower jaw is strongly denticulate,
with a well-developed median hook. The upper jaw has strong
cusps separating a median notch.
In size, alabamensis is probably similar to nelsoni. Two
mature females, one of which contained eggs ready to lay, which I
found dead on the road beside Mobile Bay on June 23, 1973, were
each almost exactly one foot in shell length. P. alabamensis is pri
marily herbivorous in the wild state. Much further work needs to
be done on this species.
Although we have considered each form of Pseudemys to be
assignable to one of four groups, relationships between members
of the genus are in fact so complicated that the categorization
given above is in some places artificial. One phenomenon that
complicates the picture is the curious parallelism that exists
between the concinna/floridana and the rubriventris sections; in
both cases northern representatives have relatively thin, low
shells, complex head patterns, ocelli on the bridge, a complex pat
tern on the plastron (at least when young), and brownish soft
parts, while southern representatives have thick, high shells, sim
plified head patterns, black smudges on the bridge, unpattemed
plastra, and intensely black soft parts. Since hybridization
between members of these two groups is rare, the parallel clinal
variations must represent similar responses to the same environ
mental selective factors. The higher and thicker shell found in
southern representatives of both sections may thus be rationalized
as a device to lessen the likelihood of being cracked by alligators,
which are only found in the south and are known to feed on turtles;
indeed many large cooters in Florida show scratches on the shell
that were probably inflicted by alligators; the success of the adap
tation is shown by the survival of the turtle. However, some of the
other differences are harder to understand; indeed, one would
have expected turtles of similar habits and general appearance
living in the same habitat to have evolved different color patterns,
for purposes of species-recognition in mating, rather than parallel
patterns.
Other curious parallelisms occur; for example, westernmost
representatives of both the concinna and floridana groups have a
cusp in the middle of the upper jaw, while Pseudemys concinna
texana shows several striking similarities with both Pseudemys
134
scripta gaigeae and Graptemys versa'. Moreover, full-species

separation of concinna and floridana is challenged by either mas
sive hybridization or complete intergradation in a number of
areas. Theoretically these phenomena differ in the fertility of the
cross-breeds; personally, I would be prepared to bet that the inter
mediates were fertile. However, in north-west peninsular Florida,
peninsularis and suwanniensis behave as perfectly good species,
living together yet rarely if ever hybridizing (though there are in
dications that intermediates occur near Tampa); yet peninsularis
intergrades with floridana, which ‘mixes’ (to use a non-committal
term) with concinna, which intergrades with suwanniensis. The
important thing is to understand the biological situation and to
realize that our nomenclatorial system, excellent though it is for
most purposes, is inadequate for situations such as this.
GENUS CHRYSEMYS
Closely related to Pseudemys are the painted turtles, Chry
semys. The skull is virtually identical to that of Pseudemys, and
some herpetologists regard the two genera as inseparable. I keep
them separate on the grounds that Pseudemys is such a hotch
potch of confusing, intergrading, generally similar forms that it
seems unnecessary to add to it a species which can always be dis
tinguished, at a glance, from any true Pseudemys. A genus, after
all, is unlike a species in having no strict biological definition, be
ing merely a convenient and apparently natural group of related
species. In keeping the genera separate, I follow the examples of
Dr. Ernest Williams, whose pronouncements on turtle matters are
always of the greatest accuracy, and of Dr. John Legler, who has
many years of field experience with the genus Pseudemys, and has
collected several thousand specimens.
Painted turtles are smaller than Pseudemys species, have
relatively smooth shells which lack a serrated posterior margin,
and there is often a middorsal stripe at some stage of develop
ment, though never a keel. The upper jaw is notched anteriorly;
the biting surfaces of the jaws are narrow.
The four painted turtles are all races of one species, Chry-
semys picta. The eastern painted turtle, Chry semys picta picta, is
a small turtle whose range extends along the eastern seaboard of
North America from Nova Scotia and central Maine south to cen
tral Georgia and Alabama. The carapace is normally between five
and six inches long, the record length being 7.1 inches. This race is
unique among North American turtles in having aligned central
and costal scutes, at least in the front part of the shell (in other tur
tles, of course, these scutes alternate). The carapace is dark olive
or dark brown in color, with light transverse seams. The plastron
is immaculate yellow, sometimes stained with red-brown ferric
135
www.Ebook777.com
oxide. The head is marked with two conspicuous yellow spots be
hind each eye, and there are yellow stripes on the side of the head
and the chin; these lines are continued on the neck, but they
change color to bright red. Bright red markings are also present
on the underside of the marginal scutes and on the skin of the limbs
and tail. The eastern painted turtle inhabits both still and moving
bodies of water, sunning frequently but diving at the slightest dis
turbance. When captured, it struggles hard and may bite pain
fully, though not dangerously. It is a very adaptable and hardy
species, adapting well to life even within city limits, and is some
times seen swimming at moderate speeds under ice. In diet it ap
pears to be mainly carnivorous, but sometimes it will eat vegeta
tion.
The eastern painted turtle, Chrysemys picta picta. Notice the close align
ment of the costals and centrals. Photo by R.J. Church.
The midland painted turtle, Chrysemys picta marginata, is

found from the Great Lakes region of Canada and the United
States south to southern Illinois, Tennessee, and extreme northern
Alabama; there is an extensive area of intergradation with C.p.
picta extending from Alabama to Vermont. It is distinguished
from the eastern race by the alternating carapace scutes, which
lack yellow margins, and by the presence of a dark figure along
the midline of the plastron (not usually extending even half way to
ward the bridge). In size and habits this race is similar to the east
ern painted turtle. Nesting has been recorded in June and July;
recorded clutches numbered from three to eleven eggs.
136
The southern painted turtle, C.p. dorsalis, has the immaculate

plastron of C.p. picta but the non-aligned carapace scutes of mar
ginata, and in addition has a very prominent orange streak down
the middle of the back, with lesser orange streaks along the mar
gins of the costals. This race is slightly smaller than the aforemen
tioned, the record length being 6 1/8 inches. The range extends
from extreme southern Illinois to north-west Alabama and along
the Mississippi to the Gulf of Mexico. There appears to be an iso
lated colony in the southeastern comer of Oklahoma. A population
of this race has also recently been reported from Caddo Lake, Har
rison County, Texas by J.P. Jones of the Fort Worth Zoo.
The western painted turtle, Chrysemys picta belli, is found in
southern Canada and the north-west United States from Wisconsin
to Vancouver, south to Kansas and Missouri, and with several dis
junct colonies in New Mexico and Arizona, with at least one in nor
thern Mexico (in one of the rivers draining out of the Bolson de
Guzman in extreme northern Chihuahua). This is the largest and
most distinctive race of painted turtle; the record length is almost
ten inches, and adults are commonly over seven inches long. The
ground color of the carapace is green, with a reticulate pattern of
yellow lines. The head and limb striping is yellow, and the plastron
bears an extensive and complex figure that sends out branches
along the seams. The ground color of the plastron is pink or red.
Nesting occurs in May and June, and clutches from 6 to 20 in num
ber have been recorded in British Columbia, from 5 to 13 in Iowa.
In the northern parts of the range, overwintering of the young in
the nest appears to be standard practice. The claws on the fore
limbs of these (and other) painted turtles are elongate and are
used for caressing the face of the female during courtship.
GENUS GRAPTEMYS
Another genus related to Pseudemys is Graptemys, which
comprises the map turtles and sawbacks of the eastern United
States. These turtles have striped necks and limbs, sometimes
beautifully marked heads, shells with serrated hind margins, and
central scutes with raised and sometimes pointed median projec
tions (hence the common name ‘sawbacks’). A plastral pattern of
the linear, seam-following type is usually present, at least in the
young. These turtles are all found in rivers and show a marked dis
inclination to walk about on land. This has had the result that most
of the species are restricted to particular river systems. Where
these systems are extensive (such as that of the Mississippi), the
overall range of the species may be very wide; but most of the spe
cies inhabit small rivers draining into the Gulf of Mexico, and the
total range of these species is highly restricted. Sexual dimor
phism is extreme in this genus; in several of the species, the fe
males may be more than twice as long as the largest males and
137
www.Ebook777.com
develop huge heads with massive mollusc-crushing jaw plates,
while the males have singularly small heads. These sexually
dimorphic forms of Graptemys show a most interesting and un
usual phenomenon in having habitat separation of the sexes;
males are usually found around snags and submerged branches
near the banks of rivers, while the much bigger females are found
in muddy areas at the bottom and are much less frequently seen
basking. This separation of both habitat and diet enables the sexes
to effectively behave as totally unrelated species for most of the
time, and they are thus enabled to be twice as abundant as they
would be if males and females were ‘competing’ for the same eco
logical niche. This separation of habitat has also had the effect that
several large museum series of certain Graptemys species consist
entirely of males and juveniles, having been hand-collected in
vegetation and trash at the edges of the rivers.
Barbour’s map turtle, Graptemys barbouri, exemplifies this
extreme sexual dimorphism. Adult females are commonly over
ten inches long, while a shell in my possession 12.8” long may well
be the largest Graptemys on record; the skull associated with this
shell is exactly three inches wide. Females develop huge, toad-like
heads with extensive crushing plates on the jaws. Males, on the
other hand, have very small, narrow heads and are not known to
exceed a carapace length of 4.65 inches. Barbour’s map turtle is
found in the Chipola, Apalachicola, and Flint Rivers in the pan
handle of Florida and southwestern Georgia. Young specimens
have very high, saw-toothed keels along the midline of the shell,
but this keel is no longer evident in old females, though tubercles
at the rear of the second and third central scutes remain. The cara
pace is olive-green with irregular wavy yellow lines on the costals
and inverted L-shaped markings on the marginals. There are yel
low-green stripes along the neck, limbs, and tail, and a pair of nar
row lines along the nape branch out to become large greenish
areas behind each eye. The plastron is unmarked in adults, but in
juveniles the transverse seams are dark-edged.
The natural sex ratio of Graptemys barbouri appears to be
about two males to one female. In many mature females the lower
jaw projects a short distance beyond the upper. Nesting takes
place in the summer months; each female produces, on average,
about 22 eggs per season, probably in more than one clutch. The
young hatch in late October. The eggs measure approximately
1.5” x 1”.
This species is still common in the Chipola River, where Carr
and Marchand collected the type series in 1941. The Apalachicola
colonies may be large, but are very difficult to assess because of
the turbidity of the water. Sam Telford tells me that he has seen
138
Female Barbour’s map turtle, Graptemys barbouri. Notice the wide head and
relatively low tubercles.
large female barbouri in salt water near the bridge across East
Bay, off the mouth of the Apalachicola.
To the west, barbouri is replaced by the related Alabama map
turtle, Graptemys pulchra. This species is found in all major
drainages entering the Gulf of Mexico from the Pearl River to the
Yellow River, including the Pascagoula, Mobile Bay, and the Es
cambia and its tributaries. However, it is absent from the Perdido
Bay and Blackwater drainages in Florida and Alabama, and ap
parently also from the Escatawpa River, a tributary of the Pasca
goula. Cagle in 1952 recorded a single specimen of barbouri sym-
139
www.Ebook777.com
patric with pulchra in the Escambia River, but this record has
since been disproved. However, in different parts of its range, pul
chra is sympatric with Graptemys nigrinoda, G. flavimaculata,
and G. oculifera.
Graptemys pulchra differs from G. barbouri in that the cara
pace of the mature female is slightly higher, and both sexes at all
ages show a black line along the vertebral ridge. G. pulchra also
has a longitudinal light bar under the chin instead of a curved bar
following the jawbones. Sexual dimorphism is extreme; males
mature in three or four years and reach a maximum length of 4.3”,
while females mature in about fourteen years and may reach a
length of 11.25”. The head of mature females becomes greatly
swollen, as does that of barbouri, but the snout of pulchra is always
more pointed, and the lower jaw does not become undershot. The
skull of pulchra is said to be unique in the genus in having the jugal
bone excluded from the orbit; this is Indeed the case in the three
skulls in my possession, although Archie Carr found the character
of the jugal and its relation to the orbit to be “unreliable.”
The carapace of G. pulchra is olive, with distinct curved yel
low markings on each of the marginal scutes and irregular, wan
dering yellow lines on the costals. The hind margin of the shell is
markedly serrate, and the nuchal area is strongly indented. The
plastron is yellow, though often stained dark red-brown, and there
are dark linear markings on the bridges and under-marginals at
all ages. In addition, young specimens have dark transverse plas-
tral seams. The head markings of the Alabama map turtle are
highly distinctive. The entire area between the orbits is occupied
by a single very pale green or almost white, anteriorly tapered
figure, and there is another large area of a similar color behind
each eye. In five young specimens from the Conecuh River, Ala
bama, in my possession, the lateral markings do not make contact
with the median one; but they frequently make at least narrow
contact in other river systems to the west. The neck and limbs are
marked with numerous pale stripes and there is a delicate
X-shaped marking behind the median pale area on the front of the
head. There appear to be differences between the populations of G.
pulchra in the Pearl River, the Pascagoula River, the Mobile Bay
drainages, and the Escambia-Yellow River drainage, though these
have not yet received full subspecific recognition. The Mobile Bay
drainage, in which the species extends above the Fall Line and is
subject to turbulent water, is characterized by turtles with unusu
ally flattened shells. In the Pascagoula, the turtles have a greyish
tinge to the green ground color and have relatively light head and
marginal scute pigmentation. Females, at least, from the Pearl
River differ from those of the Escambia River in having the mark
ings of the lower jaw tending to be broken into erratic lines that
may make up either a longitudinal or a transverse pattern.
140
Until recently the Alabama map turtle was very poorly

known; however, studies of the species by Dr. Robert Shealy of
Pensacola Junior College have added enormously to our know
ledge of the life history of pulchra, and now it is perhaps one of the
best-known species of the genus. Much of the life history data
given below was discovered by Dr. Shealy.
The Alabama map turtle feeds for six months of the year, from
April to October; during the remaining months, true hibernation
does not normally take place, but activity levels are dependent
upon prevailing temperatures. In both sexes growth slows down
markedly after maturity is reached; maximum size is reached in
males at about 8.4 years and in females at about 23 years, after
which age and size are no longer correlated. However, the skull in
the female continues to increase In width after the shell has reach
ed its maximum size. Courtship apparently takes place from Sep
tember to November, which would suggest that sperm is stored by
the female until the following nesting season. Immature speci
mens in captivity sometimes show precocious courtship activity,
which involves titillation of the head of the female with the snout.
Copulation has not been described; in view of the disparity in size
between the sexes, it should be interesting to watch. Dissection of
mature females suggests that egg production for a season varies
from 7 to 71, with an average of 29; there is a distinct correlation
between clutch size and female shell length. One to six clutches
may be laid in the course of a season. Eggs from the Conecuh
River population averaged 38 mm by 26 mm; on the other hand,
eggs from a Pearl River female measured, on average, 45 mm by
26 mm. The eggs have a leathery, finely granulated texture when
newly laid. Nesting takes place from late April to late July, typic
ally 3 to 15 meters from the water’s edge in banks of fine sand asso
ciated with sharp bends in the river. The track leading from the
water to the nest may be quite tortuous, but the return track is al
ways direct. Fish crows and raccoons destroy a large percentage
of nests—as many as 95% in the Conecuh River, about 50 to 75% in
small streams, such as Opintlocco and Cubahatchee Creeks, which
drain into Mobile Bay. Undisturbed nests hatch in 74 to 79 days.
However, once the young turtles succeed in reaching the water,
their survival prospects appear to be very good; Shealy found
yearlings to be as numerous in the spring as hatchlings had been
the previous fall.
Juveniles and males of G. pulchra feed almost entirely upon
insects; Trichoptera, Coleoptera, Odonata, Hymenoptera, and
Diplopoda have been identified in fecal samples. However, adult
females feed almost exclusively upon the mussel Corbicula manili-
ensis, an Oriental species first introduced to the western United
States in 1938 and now found clear across the country; it has reach-
141
www.Ebook777.com
ed such a density in some parts of the range of pulchra that the bot
tom substrate contains as much mussel as sand.
Throughout its range, the Alabama map turtle spends a great
deal of time basking. Basking reaches its peak at the beginning of
the season, in March and April, when water levels are high; bask
ing remains extensive from May through July but normally de
clines from July to October. Preferred basking sites are semi-sub-
merged fallen trees and branches with no connection remaining to
the bank, relatively free of smaller branches, and having an unob
structed visibility over a wide angle. Shealy found no basking on
cloudy days when water temperature exceeded air temperature
and concluded that the principal function of basking was thermo
regulatory.
The black-knobbed sawback, G. nigrinoda, is found in the Ala
bama, Tombigbee, Cahaba, Coosa, Black Warrior, Mobile, Ten-
saw, and Tallapoosa Rivers of Alabama; in Mississippi specimens
have been collected from the Tombigbee in Lowndes County. Two
subspecies exist: the nominate form in the upper reaches of the
rivers, and delticola near Mobile Bay; there is an extensive area of
intergradation. Graptemys nigrinoda has a moderately low cara
pace, while the vertebral projections, sharp in all other species,
are blunt, broadened, and knob-like. The carapace is grey-brown,
with narrow, dark-edged ocelli on the costals. The southern sub
species has an extensive, elaborate plas.tral figure composed of
transverse and oblique bars, while the northern race has a reduced
figure never occupying more than 30% of the area of the plastron.
The races also differ in the greater width of the light markings on
the soft parts in the northern subspecies. The head pattern is very
variable but usually includes short light transverse bars behind
the eyes, which connect in angular fashion with the paired light
lines along the nape. The ground color of the plastron is orange or
yellow. The carapace margin is serrate both laterally and posteri
orly in young specimens. The head is small even in big females.
The record shell length is about eight inches for females, about 4*4
inches for males; the largest specimens are found in the southern
part of the range.
The yellow-blotched sawback, Graptemys flavimaculata, is
found in the Pascagoula River system of Mississippi, where it is
locally known as the ‘bumpy back turtle.’ This is a small species;
the largest female in the type series had a carapace length of about
seven inches, and the males are, as usual, considerably smaller
than the females. This is another narrow-headed species. There is
a broad horizontal bar or blotch behind each eye and a large yellow
blotch on each of the costal scutes and on each side of each verte
bral. These blotches are highly variable in shape, ranging from
discoidal to curved and elongate, but they do not usually form
Comparison of the yellow-blotched sawback, Graptemys flavimacutata

(above) and the ringed sawback, Graptemys oculifera (below) in ventral view.
Specimen above from the Lantz collection.
143
www.Ebook777.com
complete ocelli. The vertebral spines are sharp, and the plastron,
light yellow in ground color, has a central dark figure of the seam
following type. The soft parts are pale olive-green with dark
stripes. A strong prefrontal ‘arrow’ is present. The Pascagoula
River where this species lives is highly polluted by the effluent
from sawmills and other industrial plants; however, Fred Cagle,
the describer of the species, considered it possible that this pollu
tion has in fact favored Graptemys populations by increasing mol
lusc populations, and Peter Floyd writes that even today it is possi
ble to see over 400 basking flavimaculata on a clear summer day
on the Pascagoula; the highest concentrations are between Luce-
dale and Escatawpa.
Male flavimaculata have elongated claws on the forelimbs,
and presumably they use these for courtship in a manner similar
to Pseudemys and Chrysemys. The yellow-blotched sawback nests
on sandbars or adjacent high ground from May through August.
Typically six eggs are laid, but recorded clutches have numbered
as many as eight. The hatchlings are 1 1/8 inch to l.5 inches in
length.
The natural diet of flavimaculata is known to include a high
percentage of snails, and this is also their preferred diet under
captive conditions.
The ringed sawback turtle, Graptemys Oculifera, is limited to
the Pearl River system of southern Mississippi and adjacent Lou
isiana. It may be considered as a version of the yellow-blotched
sawback in which the yellow carapace markings, instead of being
irregular blotches, form perfect and very striking orange-yellow
ocelli on the costal scutes; incomplete but equally bold ocelli are
on the marginal scutes, while the centrals are marked with irregu
lar yellow lines. The plastron is orange-yellow with dark trans
verse seam-edges. The ground color of the soft parts is grey-black,
with pale yellow-green stripes. The stripes on the head are broad
and very vivid. The maximum length of females is 8.5 inches, of
males about 4 inches. The diet consists mostly of insects. The eggs
are markedly elongate, being about 1.6 inches long and 0.8 inch
wide. Three or four eggs are laid at a time. Females mature at a
shell length of about five inches.
We see, then, that in each of the river systems between ex
treme eastern Louisiana and extreme western Florida we have
two Graptemys types co-existing: a broad-headed form (pulchra
or barbouri), together with a narrow-headed form (nigrinoda, fla
vimaculata, or oculifera). Despite the spatial co-existence of tur
tles of each type in one river system, there is presumably ecologi
cal separation; the large-headed forms have greatly developed
jaw muscles and crushing plates, and feed on hard-shelled mol
luscs, while the narrow-headed forms lack the equipment to cope
144
with such a diet, and feed on soft-bodied organisms. This division

into broad and narrow-headed forms is maintained in and beyond
the Mississippi drainage; the common map turtle (Graptemys
geographica) and the Mississippi map turtle (Graptemys kohni)
develop broad heads, while the false and Texas map turtles (Grap
temys pseudogeographica and G. versa) always have narrow
heads; and again there is little co-existence of different broad, or
different narrow-headed species, although the common and Mis
sissippi map turtles do overlap in Missouri and northern Arkansas.
The common and Mississippi map turtles have so much
greater and more diverse ranges than the Gulf Coast broad-head
ed species (G. barbouri and G. pulchra) that there is less emphasis
on rigid specialization, and the heads even of old females do not
reach the same degree of grotesque enlargement shown by the
highly restricted and specialized Gulf Coast species.
The common map turtle is found in Lakes George and Champ
lain, westward through the St. Lawrence and the Great Lakes
drainage to Wisconsin; also the Mississippi drainage from south
ern Minnesota south to Arkansas and northern Alabama; and the
Susquehanna drainage from New York to Maryland. This species
barely shows the vertebral spines typical of the genus, and the
carapace is relatively flattened. Young specimens are very decor
ative, having complex map-like patterns on the carapace and soft
parts. The ground color of the shell is green or olive, and there is
an isolated small yellow spot behind each eye, surrounded by a
complex pattern of lines. Large females reach a length of up to
eleven inches, but males are not known to exceed 5.5 inches. Nest
ing takes place from late May to July, and clutches ranging in
number from ten to sixteen have been recorded. In diet the com
mon map turtle is entirely carnivorous, molluscs and crayfish be
ing especially favored.
The collective range of the three subspecies of the false map
turtle, Graptemys pseudogeographica, extends from southern
Minnesota and Wisconsin south to the Sabine River, which sepa
rates Louisiana from Texas. The carapace is grey-brown in color
and has flat sides and a high keel, strikingly tuberculate in the
young but less so in adults. The plastron bears a figure of the lin
ear, seam-following type in the young, but this becomes vague in
adults. The head is very small, even in adult females.
The nominate subspecies, G.p. pseudogeographica, exists in
the pure form only in southern Minnesota and Wisconsin. This race
has narrow postorbital lines and no enlarged spots on the man
dibles. This is the largest of the three subspecies, females reaching
a length of nine inches, males about half that. The area of intergra
dation with G.p. ouachitensis extends almost throughout Illinois
west to eastern Nebraska. The range of the Ouachita map turtle it-
145
www.Ebook777.com
Two subspecies of the false map turtle. Above, Graptemys pseudogeographi-
ca ouachitensis, from the central United States. Below, Graptemys pseudo-
geographica sabinensis, from the Texas-Louisiana border.
146
self extends from eastern Tennessee west to eastern Oklahoma

and Kansas, and south to northern Louisiana. This race has a
squarish or rectangular yellow blotch behind the eye, prominent
yellow spots above and below the angle of the jaw on each side, and
between one and three of the longitudinal neck lines reaching each
eye. The Ouachita map turtle, in Illinois at least, is a late nester,
the earliest nesting date on definite record being July 6. Recorded
egg complements range from seven to thirteen. In diet this species
appears to be largely vegetarian when adult, in sharp contrast to
the carnivorous common map turtle.
In the Sabine River, the Ouachita map turtle is replaced by the
Sabine map turtle, G.p. sabinensis. This race has an oval or elon
gate postorbital spot, transverse bands under the chin, between
five and nine of the light neck stripes reaching each eye, and a
plastral pattern which is distinct only In juveniles and consists of a
greater number of finer lines than in ouachitensis.
The Mississippi map turtle, Graptemys kohni, is similar in
many ways to the false map turtle, of which it was once considered
The Mississippi map turtle, Graptemys kohni, showing the distinctive cres
cent behind the eye.
147
www.Ebook777.com
a subspecies; however it can always be distinguished by the light
crescent behind each eye, which cuts off all of the horizontal neck
stripes from reaching the eye. This species is found in the lower
and western parts of the Mississippi drainage, in Louisiana,
Arkansas, and Missouri, extending to eastern Kansas, Oklahoma,
and Texas. The Mississippi map turtle can be recognized by the
distinctive eye, the iris of which has the appearance of a pure
white, unbroken ring. In the false map turtle, the iris is also white,
but there is a horizontal dark bar running through the pupil. The
young have complex linear plastral patterns, high, black-spiked
keels, and posteriorly serrate carapaces. The head of adult female
kohni is somewhat intermediate in breadth between that of the
almost microcephalic false map turtle and the broad-headed com
mon map turtle. Its dietary preferences, too, appear to be inter
mediate, both vegetable and animal matter being eaten. These
more generalized feeding habits apparently make it possible for
the Mississippi map turtle to co-exist both with the common map
(in Missouri and northern Arkansas) and with the false and Sabine
map turtles (almost throughout their ranges). It is possible that
hybridization with the false map turtle occurs on occasions.
The Texas map turtle, Graptemys versa, was also once con
sidered a subspecies of the false map, but in fact it is a very dis
tinctive form, intergrading with no other taxa and showing almost
as much resemblance to the Texas slider, Pseudemys concinna
Graptemys versa, the

Texas map turtle, from
the Colorado River
system. Specimen from
the Lantz collection.
148
texana, as to the other Graptemys. This species is restricted to the

Colorado River system in Texas. The carapace is relatively low
and the scutes have a slightly bulbous appearance. The dorsal pat
tern is intricate, consisting of fine light lines and ocelli on an olive
background. The vertebral spines are not prominent and lack the
black tips shown by most other map turtles. The plastron is im
maculate even in quite young specimens, but there is a pattern of
longitudinal lines on the bridge. This is probably the smallest spe
cies of the genus, the maximum size even for females probably
being less than six inches.
A new species of map turtle, Graptemys caglei, was first intro
duced to the herpetological world by the well-known Indiana rep
tile dealer Larry Lantz in his price list dated May 5, 1972. The spe
cies was more formally described by Haynes and McKown
(Tulane Studies in Zoology and Botany, Vol. 18, No. 4; Jan. 24,
1974). It extends the known range of the genus about 100 miles
south, being known from the Guadalupe River of south-eastern
Texas and its tributaries, the Blanco River and the San Marcos
River ; there are also sight records for the San Antonio and Medina
Rivers. Records exist for DeWitt, Gonzales, Hays, and Kerr Coun
ties, Texas. This is a small species, the only adult female in the
type series having a carapace length of 6.35 inches, though there
are sight records of larger specimens. A typical mature male
measures about 3.5 inches in carapace length.
In the opinion of Haynes and McKown, caglei is a morphologi
cal intermediate between G. versa and G. kohni, which are nar
row-headed and broad-headed species respectively. G. caglei is
immediately distinguishable by the delicately marked, predomin
antly green carapace, that of versa being olive and of kohni brown
ish with black markings. The carapace, being flattened and knob
by, is more similar to that of versa than kohni. The vertebral
spines, which are marked with brown or black, are of intermediate
length. The front part of the head, in lateral profile, is horizontal,
as in kohni, rather than downsloped as in versa. The head pattern
is rather similar to that of kohni, having a strong V-shaped mark
on the top, the sides of which curve down behind each eye to form a
crescent. However, whereas the light postorbital mark is single in
kohni, in caglei there are one or two parallel markings behind of
almost equal intensity. The plastral pattern of caglei is of the typi
cal type, with dark lines closely paralleling the ventral seams,
though in some individuals there are also numerous small black
flecks on the plastral scutes. The ground color of the soft parts is
black, with numerous thin cream-colored lines, including seven on
the dorsal surface of the head and approximately twenty-two along
the neck. The limbs have a well-defined pattern of broad and nar
row cream-colored stripes.
149
www.Ebook777.com
Comparatively few biological data on this species are avail
able as yet. However, caglei is reported to be abundant in various
sections of the Guadalupe River and the San Antonio River, where
in places it may be the dominant-chelonian species. Nesting has
not been described, and since sandbars are essentially absent in
many parts of the Guadalupe River, the nesting habits may be dif
ferent from those of other Graptemys from the Gulf Coast drain
ages.
From stomach analyses of juvenile specimens, both plant and
animal matter may be consumed, though it is probable that most
of the plant matter found (bark, algae, and grass fragments) may
have been accidentally ingested. Large quantities of very small
dipteran insects found in the stomachs suggest that the turtles
may have fed upon deposits of already dead insects that had accu
mulated against twigs or other items at the water’s surface. The
adult males apparently feed largely upon caddis fly larvae, while
subadult females are known to eat small, thin-shelled gastropods.
The three genera we have d. : so far—Pseudemys, Chry-
semys, and Graptemys—are clearly closely related to each other.
Several superficial features confirm this relationship—the appear
ance of similar linear plastral patterns in all three genera, the
neck and limb striping, the horizontal bar through the eye, and so
on; moreover, while these color pattern similarities could concei
vably have evolved independently, one peculiar structural and be
havioral feature common to the three genera is so bizarre that it
surely confirms their common origin. This feature is the courtship
pattern, in which the male strokes the face of the female with his
sometimes absurdly elongated fingernails; this has been observed
in many Pseudemys species, in Chrysemys, and in Graptemys, at
least in G. flavimaculata.
GENUS MALACLEMYS
Closely related to Graptemys is Malaclemys terrapin, the dia-
mondback terrapin, which shares the Graptemys features of con
siderable sexual dimorphism, enlarged heads in adult females,
and vertebral knobs somewhat similar to those of Graptemys nig-
rinoda. The diamondback terrapin, of which seven subspecies are
recognized, is the only American emydid turtle adapted for life in
brackish and salt water; this species is found in salt marshes, es
tuaries, and protected lagoons from Massachusetts along the en
tire Atlantic and Gulf coasts of the United States. This is a medi
um-sized species, adult females being six or seven inches long and
males four or five inches. Differences between Malaclemys and
Graptemys are obvious, though superficial; Malaclemys has a
spotted instead of a striped pattern on the soft parts, usually has
quite strong concentric annuli on the carapace scutes, and the hind
150
A large female diamondback terrapin, Malaclemys terrapin. Photo by Dr.

Herbert R. Axelrod.
margin of the carapace typically curls up instead of flaring. The

hind legs of diamondback terrapins are large and the toes have ex
tensive webs; these are vigorous turtles, swimming fast and strug
gling actively when picked up. The flesh is considered more succu
lent than that of any other turtle, and from the late nineteenth cen
tury until about 1920 diamondback terrapin was a very expensive
and much sought-after gourmet item. Only the northern subspe
cies were considered really good to eat, and large farms were es
tablished (notably at Beaufort, North Carolina) to breed the spe
cies in captivity. After the First World War, people seemed to lose
interest in terrapin, possibly because of the depression and prohi
bition (terrapin being properly prepared with liquors of various
sorts), but more likely, as Archie Carr suggests, because diamond-
back, though very good, is not whole orders of magnitude better in
taste than many much cheaper turtle species, and the vogue for
terrapin, as opposed to snapper or soft-shell, was really a partially
irrational fad. Roger Conant suggests that another reason for the
loss of interest in terrapin was the decreasing availability and in
creasing cost of household help, a terrapin dinner taking many
hours to prepare properly.
The northern race of the terrapin, Malaclemys terrapin terra
pin, is found from the northern shore of Cape Cod south to Cape
Hatteras, North Carolina, where intergradation with M.t. centrata
151
www.Ebook777.com
Hatchling diamondback terrapin, Malaclemys terrapin, showing the unusual
tubercles of young specimens. Photos by R.J. Church.
occurs. These two races are very similar, but terrapin tends to
have a more wedge-shaped carapace than centrata (when viewed
from above) and a more strongly sculptured pattern of annuli on
the carapace. The spotted pattern of the head and neck is very var
iable. Terrapin dealers of the Gay Nineties recognized many local
varieties of terrapin by their color pattern, but the habit of ship
ping large consignments of terrapins from one place to another
has made it impossible now to assign particular color varieties to
particular localities.
These two northern subspecies appear to have much the same
habits and habitat preferences. Animal matter (particularly crus
taceans and molluscs) predominates in the diet, but on occasions
tender shoots and rootlets are eaten. In captivity they thrive on
chopped fish. These terrapins hibernate in mud and mate immedi
ately upon waking up in the spring. Eggs are laid in May and June,
and clutches number from five to twelve eggs. Hatchlings are just
over an inch in length. Large basking aggregations of these terra
pins are sometimes seen on warm days.
Malaclemys terrapin centrata reaches as far south as Volusia
County, Florida, where it intergrades with the Florida east coast
terrapin, Malaclemys terrapin tequesta, which has a dark and al
most smooth carapace. This race was described in 1955 from spe
cimens from Miami Beach, Florida; its habits have not been
studied.
Diamondback terrapins from the lower Gulf Coast of Florida
south of Fort Myers, and from the Florida Keys and the
Marquesas, are commonly referred to as the subspecies
rhizophorarum. Supposed characters of this subspecies mentioned
by Archie Carr include the strongly oblong shell and the bands of
152
black pigment along the seams of the lower marginals and

frequently the plastral scutes as well. Nevertheless, this race has
still not been properly characterized, and it appears to be
inexplicably rare or absent from extensive sections of the range
commonly ascribed to it. Terrapins are not uncommon in Florida
Bay, but most of the individuals on the mainland side have the
light scute centers typical of the race macrospilota and
presumably represent an intergrade population. In the Keys
themselves, terrapins have been found on Key Largo (at the
extreme upper end of the chain of islands) and on Key West, from
Barracuda Key (just north of Key West), the Marquesas Islands
(between Key West and the Dry Tortugas), and Boca Grande Key,
between Key West and the Marquesas. Between the Upper Keys
and Lower Keys populations, however, lies a hundred miles of
islands from which terrapins are apparently unreported, despite
what looks like a lot of suitable habitat. Moreover, the turtles from
the Upper and Lower Keys are apparently dissimilar, though the
available sample size is too small to be sure. The Key Largo turtles
may typically show the vertical striped pattern on the rump that
Conant considers to be characteristic of rhizophorarum, and old
females may have exceptionally large dark spots on the head and
limbs. A Key Largo juvenile described by Schwartz had bold,
broad, doughnut shaped figures on the coastal scutes (hollow on
coastals 1 to 3 and solid on coastal 4) and one to four dark spots on
each of the plastral scutes.
The turtles from the Lower Keys apparently lack the vertical
striping on the rump, judging by a series of eight live specimens
collected by Gale Belinky, and have uniformly dark shells with
markedly expanded dorsal tuberosities. The juveniles have
freckles on the skin, but old females may have large spots on the
head that may fuse into irregular streaks. However, larger
numbers of individuals of both populations must be examined
before the range of variation is known, since diamondbacks of
other subspecies are notoriously variable in skin pattern even
within a single population.
The habits of rhizophorarum (which may in fact be a
composite of two subspecies) have not been studied
systematically. However, the subspecific name is apt, the turtles
usually being found in areas of dense red mangrove (Rhizophora),
though Carr was informed by several people that the terrapins are
sometimes found in spring-tide and hurricane pools inland on dry
keys and isolated from permanent water. One of the large Key
Largo females kept by Gale Belinky excreted fragments of
hundreds of mussels (Mytilus recurvus). Maximum shell length
appears to be about 8.25 inches.
153
www.Ebook777.com
The ornate diamondback, Malaclemys terrapin macrospilota,
is easily recognizable by the orange or yellow centers of the cara
pace scutes. The central keel is tuberculate, as in rhizophorarum.
Hatchlings have uniformly horn-colored shells, but the tubercles
on the second, third, and fourth vertebrals are very striking. This
turtle nests in April to June, and egg counts range from four to ten.
Nesting usually takes place in or around sand dunes, sometimes a
long way from the water. There is some evidence that large num
bers of females nest synchronously in Tampa Bay, but the implica
tions of this observation have not been investigated. The ornate
diamondback ranges from the southern part of the Gulf Coast of
Florida to the Panhandle; at the northern limit of the red man
grove, intergradation with the Mississippi diamondback takes
place. A very beautiful phase of the ornate diamondback which is
occasionally found near Tampa has a pure white head with no dark
spots.
The Mississippi diamondback, or Biloxi terrapin, M.t. pileata,
ranges from the Florida panhandle to eastern Louisiana. This is
the largest of the races, a 9.37 inch female being on record. It is
very similar to the mangrove terrapin, but it lacks the black plas-
tral seams of that race. The central keel is tuberculate, and the
upper lip and the upper surfaces of the head are usually dusky. The
hind margin of the carapace is strongly curled upward. Old fe
males get exceedingly broad heads and have rather lighter lips
than the males.
The Texas diamondback, Malaclemys terrapin littoralis, in
tergrades with the Mississippi race in eastern Louisiana, and its
range extends to the vicinity of Corpus Christi, Texas in the south.
Records from the Yucatan Peninsula, however, are based on
misidentifications of Rhinoclemmys areolata. The Texas diamond-
back has a tuberculate keel in young individuals which is lost in old
adults. The shell is higher and the bridge deeper (in lateral view)
than in other races, and the highest point of the carapace is further
back. The upper lip is white, not infused with black as that of the
Mississippi race. The habits of this race have not been studied.
GENUS DEIROCHELYS
A turtle whose superficial similarities to Pseudemys and
Chrysemys may or may not indicate actual close relationship to
those genera is the chicken turtle, Deirochelys reticularia. This is
a medium-sized species, commonly six or seven inches long, the
record being about ten inches. The shell is long and narrow, quite
low in small specimens but steeply domed in big females. The plas
tron is large and is immaculate yellow in color in all except the
westernmost subspecies, while the bridge has a variable degree of
black pigmentation. The carapace is marked with irregular
154
reticulations, and the entire surface is sculptured with fine longitu

dinal wrinkles. The hind margin is smooth. The soft parts are
striped, the most prominent head stripe extending backward and
downward from the corner of the eye. There is a broad longitudinal
yellow stripe on the front face of each of the forelimbs, with a nar
rower line above it. The skin just in front of the axillary notch, and
between the hind legs and the tail, is marked with vertical yellow
bars. The physiognomy is entirely different from that of Pseu-
demys, the head being rather flat and the eyes close together. The
neck is exceedingly long, and the retractor and extensor muscles
are housed under the specialized dorsal rib-ends, which loop away
from the shell leaving a longitudinal channel on each side of the
backbone. This channel between the ribs and the shell is found in
turtles which make sudden snapping movements of the neck, such
as the American snapping turtles and the South American mata-
mata. The chicken turtle apparently shows several adaptations
which assist it in grabbing and swallowing active prey of consider
able size—probably mostly crayfish; the mouth is very large, the
disposition frequently highly aggressive, the neck extremely long
and extensible, and the throat very capacious. The jaws, however,
are relatively weak and have a simple cutting edge, with none of
the complex alveolar ridges of Pseudemys or the crushing plates
of Graptemys.
Plastral view of the

western chicken turtle,
Deirochelys reticularia
miaria. Specimen from
www.Ebook777.com
The chicken turtle occupies a wide variety of aquatic and
semi-aquatic habitats, principally ditches and shallow ponds. It
has the thinnest shell of all the Floridian emydids, and this may re
flect the fact that it is not typically found in bodies of water large
enough to house alligators. It is sometimes seen basking, but never
in large aggregations. Chicken turtles are very given to walking
about on land, and frequently get run over on highways. In capti
vity they do well enough, but are very liable to bacterial erosion of
the shell—more so than Pseudemys and Graptemys.
The chicken turtle derives its vernacular name from the suc
culence of its flesh; formerly this was a favorite eating turtle
throughout the south, but I have not heard of it being eaten nowa
days. It never became a national fad on the scale of the diamond
terrapin, probably because it was not procurable in quantities, but
the reasons for its decline in favor are not obvious.
The Florida chicken turtle, Deirochelys reticularia chrysea,
has orange or yellow reticulations on the carapace which are most
obvious in young specimens. The plastron is immaculate, and
while there are reduced markings on the bridge, the undersides of
the marginal scutes are usually unpigmented. The carapace has a
prominent orange or yellow rim. This race is only found in the pen
insula of Florida. The eastern chicken turtle, D.r. reticularia, is
duller in color than the Florida race, has narrow greenish or
brownish carapace reticulations, and has extensive black mark
ings on the bridges as well as prominent black blotches on the
undersides of the lateral and posterior marginals. This race is
found along the coastal plain from eastern North Carolina to the
Mississippi River in southern Louisiana. The western chicken tur
tle, Deirochelys reticularia miaria, found in Louisiana and parts of
Texas, Oklahoma, and Arkansas, has broad but poorly contrasting
reticulations on the carapace, no markings on the underside of the
neck in adults, and dark markings along the plastral seams.
The breeding habits of the chicken turtle have not been studied
in detail; however, in Florida nesting takes place in any month of
the year, while in South Carolina nesting does not occur in winter
but may commence as early as mid-March. Clutches numbering
between five and fifteen eggs have been recorded. The nest is cy
lindrical, about four inches deep and three inches in diameter at
the mouth. The eggs have tough but flexible shells and are about
1.5 inches in length and slightly less than an inch in width. There is
some evidence that several clutches may be laid per year. A nine-
inch female chicken turtle which I found freshly shot near my
home in Gainesville, Florida, on January 24, 1972, contained ten
eggs ready for laying, ten ovarian yolks about inch in diameter,
and numerous very small ovules. The hatchlings, which may over
winter in the nest, are almost circular and measure a little more
than an inch in length.
156
The chicken turtle is not found in moving water, and probably

for this reason the subspecies miaria and reticularia are separated
by the Mississippi River. Chicken turtles often become thickly
coated with algal growths in the wild state, but since they shed the
outer layers of the scutes as they grow (as do Chrysemys and
Pseudemys), the entire algal growth is periodically shed also.
GENUS EMYDOIDEA
Blanding’s turtle, Emydoidea blandingi, is a northern species
that used to be considered congeneric with the European pond tur
tle, Emys orbicularis, but modern opinion places it closer to Deiro
chelys than to any other form. Blanding’s turtle is found in the
Great Lakes region of the northern United States and southern
Canada; the range extends from north-eastern Nebraska to On
tario and northern Ohio. The species has apparently retreated in
modern times, as is suggested by the discontinuous distribution in
Illinois and the presence of an isolated colony in New Hampshire
and Massachusetts. Blanding’s turtle has the general shape and
size of a chicken turtle; the shell is elongate, moderately elevated
but rather flat-topped, and typically is seven or eight inches long.
Plastral view of Blanding’s turtle, Emydoidea blandingi, of the northern

United States and adjacent Canada.
www.Ebook777.com
The record length is 10.25 inches. The plastron is large, flat in the
female and slightly concave in the male, and has a well-defined
hinge between the pectoral and abdominal scutes. However, clos
ure of the shell Is not nearly as complete as in the box turtles. The
carapace is horn-colored, thickly speckled with black. The plas
tron is yellow, but the juvenile (postero-lateral) area of each plas-
tral scute is black. The dorsal side of the neck is speckled, while in
front of the eyes these black spots coalesce to form a uniformly
black area. The lower jaw, chin, and throat are bright yellow. The
eyes are red-brown. The skull and jaws are amazingly similar to
those of the chicken turtle, but the neck is not as long and the dispo
sition is much more placid. The bridge is much more shallow than
that of Deirochelys, and the gular region is planar rather than con
vex.
Blanding’s turtle is a semi-aquatic species, being found about
as frequently in ponds, small streams, and the edges of lakes as on
land. Like certain other northern species, it is highly resistant to
cold and is sometimes active when air temperatures are below
freezing. On the other hand, the critical thermal maximum of this
species is about 39.5° C., considerably lower than that of most
emydids. Copulation takes place in the water and has been record
ed from March to October. Nesting takes place in June and July,
and clutches numbering from six to eleven eggs have been record
ed. The nest is flask-shaped, about seven inches deep and 3 to 4
inches in diameter at the mouth. The eggs measure approximately
1.4” x 1”. Two clutches may be laid in a season.
This species is almost entirely carnivorous, crustaceans being
the favorite food, followed by insects.
GENUS CLEMMYS
Two other North American emydid genera, Clemmys and Ter-
rapene, continue the terrestrial trend begun by Emydoidea. Clem
mys,, as this genus is now understood, consists of four North
American species, three in the north-east and one in the south
west. Clemmys insculpta, the wood turtle, is found in Nova Scotia
and throughout New England, south to the northern tip of Virginia;
to the west it is found in southern Ontario, northern Wisconsin, and
north-eastern Iowa. It is found throughout Pennsylvania, but has
not been recorded from Ohio or Indiana. The wood turtle is readily
identified by the very rough, keeled, rather flat shell, the orange
color of the throat and limbs, and the often jet black head. The tail
is long—in the hatchlings about as long as the shell. The adult
length is about six or seven inches, the record being nine inches.
This is a terrestrial species, showing up near water only for hiber
nation or mating purposes. It has a wide tolerance of different hab
itats, and this adaptability makes it a very hardy pet animal.
Many pet keepers have remarked on the responsiveness and intel-
158
The wood turtle, Clemmys insculpta, a species now protected over much of
its eastern United States range. Photo by R.J. Church.
ligence of the wood turtle, and it does indeed appear to have

superior ability at following mazes and in reward-association ex
periments. In diet the wood turtle is truly omnivorous. Nesting
takes place in May and June, and recorded clutches range from
four to twelve in number. This species is protected by law in New
York State, but unfortunately protective laws cannot lessen the
large numbers of wood turtles killed by automobiles each year.
The spotted turtle, Clemmys guttata, is much smaller than the
wood turtle, the record length being five inches and the normal
adult length about four. The plastron is similar in color to that of
the wood and Blanding’s turtles—basically yellow, with black
areas extending from the juvenile part of each scute. The cara
pace, however, is smooth, black, and decorated with yellow spots.
Young specimens have relatively few spots—sometimes none at
all—but as the animal grows more spots develop on the costal and
central scutes. The head and limbs too are black with yellow spots.
The jaws are whitish, and frequently there are elongate orange
markings near the comers of the mouth.
159
www.Ebook777.com
The spotted turtle is found from extreme north-eastern Illinois
east to southern Maine and Maryland, and the range extends down
the coastal plain to South Carolina. Spotted turtles have also been
found at Riceboro, Georgia, and in Alachua, Marion, Suwannee,
Lafayette, and Duval Counties, Florida. However, the total
number of specimens collected in Florida is barely a dozen, des
pite the high density of herpetologists in the area, and there is
some question that the species is truly native to the State. The
spotted turtle copulates either on land or in the water, usually in
April, and eggs are laid in June. The eggs are only slightly smaller
than those of the much larger wood turtle, but are much fewer in
number—recorded clutches numbered between one and four eggs,
usually three. Conant found spotted turtles much easier to see and
catch in the spring than at any other time of the year. In diet this
species is predominantly carnivorous.
Closely related to the spotted turtle, and similar in general
form and much of its range, is the bog turtle, Clemmys muhlen-
bergi. This may be the smallest species of turtle in the world; the
record carapace length is only 4.25 inches. Moreover, American
musk turtles that barely exceed this length have deeper shells and
much bigger heads. The bog turtle has a dark brown shell with
vague reddish areas visible under strong light. The soft parts are
dark brown, with red edges to the limb scales. The plastron is
mainly dark brown to black, but has roughly symmetrical unpig-
mented areas around the midline, especially towards the front.
The bridge is dark, and there are triangular dark markings along
the under surface of the marginals. The most characteristic fea
ture of this attractive species is the large, irregular, and highly
variable area of orange, red, or yellow, or occasionally pink, loca
ted on the temporal and tympanic region on each side of the head.
The bog turtle is indeed a bog dweller, and it has perhaps the
most rigid habitat preferences of any North American freshwater
turtle. It is usually found in sphagnum bogs, in situations where it
can walk about in very shallow water. It has been said that the
average bog turtle in the wild has its feet wet and the middle of its
carapace dry. However, in the southern parts of its range it is often
found in shallow streams.
The range of the bog turtle shows two levels of discontinuity.
In certain areas where there are many bogs, each individual bog
may have its own isolated population of muhlenbergi, separated
from neighboring populations by anything from a few hundred
yards to several miles. Moreover, the overall range of the species,
as plotted on a small-scale map, shows discontinuities of dozens or
even hundreds of miles. The overall range of the species may be
summarized as follows:
160
Clemmys muhlenbergi, the bog turtle. This species is most abundant in open,
sub-climax ecosystems and may be less threatened than previously thought.
Photo by R.J. Church.
The principal home of muhlenbergi is a block of land extend

ing from southern New York State (Duchess and Ulster Counties),
east to Berkshire County, Massachusetts, and Litchfield County,
Connecticut, and south through most of New Jersey and south
eastern Pennsylvania, reaching as far south as Carroll, Baltimore,
Harford, and Cecil Counties, Maryland, and Cape May County,
New Jersey, and west as far as Franklin County, Pennsylvania.
Much of this area is of course highly urbanized, and consequently
the bog turtle colonies are quite scattered. A once abundant colony
on Staten Island, from which Raymond Ditmars obtained most of
his specimens, has now disappeared.
In addition to the range outlined above, there are several out
lying colonies of bog turtles in New York State, in Albany and War
ren Counties, in the Finger Lakes region, and in Genessee County.
At one time there was a colony in extreme north-western Pennsyl
vania, in Crawford and Mercer Counties, but this colony appears
to have been wiped out by the construction of the Pymatuning
Reservoir, nearly forty years ago.
Bog turtles are also found in the area of western North Caro
lina, from Clay to Allegany Counties, located at altitudes of up to
4000 feet and separated by hundreds of miles from the northern
populations. Investigations by Ken Nemuras have recently extend
ed the limits of this southern part of the range into at least three
counties of Virginia (Floyd, Patrick, and Carroll Counties), and
outlying colonies are known in Mecklenberg, Iredell, and Forsyth
Counties, North Carolina; but the old record from extreme north-
161
www.Ebook777.com
ern Virginia appears to have been based on a misidentified spotted
turtle, according to Jim Weaver.
Even in apparently suitable habitat within the overall range,
bog turtles sometimes prove impossible to find, and the impres
sion is of a species undergoing rapid retreat—accelerated in re
cent years by swamp drainage and urbanization of many parts of
its range. The bog turtle has unusually high thermal preferences
for a northern turtle; ideal habitat includes high tussocks of grass
which the turtle can climb on to bask, with half an inch or so of cool
water running below. Despite the spotty distribution, in suitable
habitat populations as high as 50 or more per acre have been re
corded. A virtual cult of the bog turtle has grown up among some
north-eastern herpetologists, who know of undisturbed colonies of
bog turtles which they visit from time to time, but of which neither
love nor money would force them to divulge the whereabouts.
The different colonies of the bog turtle show slight average dif
ferences in mature size and coloration, as well as in average clutch
size. In some areas three eggs are the rule, while in others clutches
of up to five eggs are frequently found. These turtles do well in cap
tivity, and some serious hobbyists have managed to breed them
through several generations. Copulation takes place in May and
early June, and most nesting takes place in June and July. The
eggs, which measure about 1.2” x 0.6”, are buried outside the bogs
in dry ground. Hatchlings are about an inch long and about 0.8”
wide.
The bog turtle is an omnivorous species; recorded dietary
items include insects, berries, seeds of pondweed and sedges (Pot-
amogeton and Car ex respectively), young snails (Succinea), mil
lipedes, and carrion, including frogs.
The fourth species is Clemmys marmorata, the Pacific pond
turtle. This turtle is somewhat similar in appearance to the Euro
pean pond turtle and achieves about the same adult size (around
six inches), though the head is smaller. The dorsal surfaces are
olive in color, speckled with black; the plastron is yellow, often
with dark markings along the rear margins of the scutes. There is
a dark bar through the eye. The tail is long, with the vent, as usual,
located more posteriorly in males than in females. Males also have
a more depressed shell than females. Two subspecies are recog
nized— C.m. marmorata from southern Vancouver and British
Colombia south to central California, and C.m. pallida in southern
California and northern Baja California. Isolated colonies of C.m.
marmorata exist in Washington State and western Nevada. The
northern subspecies differs from the southern is having inguinal
scutes (in most individuals) and in having the throat lighter in
color than the sides of the head. The Pacific pond turtle is thor
oughly aquatic and is only found away from water during the nest-
162
ing season. It is found in ponds, lakes, and streams, but prefers

quiet, muddy water. In diet it is carnivorous by preference, but
some individuals have been seen eating lily pads. Population num
bers, particularly of the southern subspecies, have been substan
tially reduced from primordial levels, partly by high demand for
food until comparatively recently, partly by habitat destruction.
Nesting of the Pacific pond turtle takes place in June and July,
with smaller numbers nesting in late April, May, and August. Re
corded clutch sizes range from three to eleven eggs; the eggs are
somewhat variable in size, being 1.3” to 1.7” in length, 0.8” to 0.9”
in width. The hatchling is about one inch long, with a tail almost as
long as the shell.
GENUS TERRAPENE
The two remaining New World emydid genera, Terrapene and
Rhinoclemys both include some completely terrestrial species
which show a number of parallelisms with the true tortoises of the
family Testudinidae. The familiar American box turtles constitute
the genus Terrapene. A variety of races exist which have now been
grouped into four species. Two of these are plentiful and wide
spread in the United States; all four species are found in Mexico,
but they are not conspicuous members of the Mexican fauna, and
the extant populations in that country are isolated and quite poorly
known.
The box turtles are easily recognized by their relatively
domed shells, well-developed plastral hinges which enable the ani
mals to close themselves off completely from the outside world,
their small size, terrestrial habitat, and neither webbed nor ele
phantine feet. The plastral hinge, as in all the hinged emydids, is
situated between the pectoral and abdominal scutes and between
the hyoplastral and hypoplastral bones.
The common or eastern box turtle, Terrapene Carolina Caro
lina, is found from the southern tip of Maine through all except
northern Pennsylvania west to the southern half of Illinois, and
south to northern Mississippi, central Alabama, and southern
Georgia. The carapace is usually rather domed, with a longitudi
nal keel. The color pattern is extremely variable, but usually the
carapace is decorated with irregular or elongate yellow or orange
smudges on a dark brown or black background. The plastron may
be any color from uniform very dark brown to pale yellow. Males
are readily distinguished by their bright red eyes and their plas
tral concavity and are usually slightly larger than females. The
head and limb markings are as variable as the shell pattern, but
typically the head and neck are predominantly yellow on a dark
background. Hatchlings are much flatter than adults, do not have
a functional plastral hinge, and often have a single yellow spot on
163
www.Ebook777.com
each of the vertebral and costal scutes. The upper jaw of adult spe
cimens is strongly beaked. The normal adult length of the common
box turtle is between four and five inches, but a recently discover
ed specimen from Glendale, New York, had a rather flattened,
bilobed carapace 7 13/16” in length.
The common box turtle is found in open woodlands, frequently
but by no means invariably in the general vicinity of water. They
are poor swimmers, floating very high, and they rarely enter
water more than an inch or so in depth. Box turtles are as variable
in disposition as they are in coloration; they never bite when pick
ed up, but some individuals struggle and show no signs of fear
when handled, while others may close up tight and not appear
again for hours.
The common box turtle, despite its small size, seems to be
among the longest-lived of turtles; there are numerous records of
specimens reaching ages of 40 to 50 years, and the following quota
tion would indicate that a much greater age is sometimes reached.
A specimen of T. Carolina found in 1931 was described as
“A very ancient female. Her carapace was absolutely
smooth, the edges so worn that there was no flare whatever.
She had been marked with a date in a rather unusual place,
across the anterior half of the plastron. The date was 1809—
nearly worn away, but I think there is no doubt about it, and
judging from the extreme age of the turtle, I believe it to be a
true date. ”
This is, of course, no conclusive proof that the turtle was, in
fact, more than 123 years old, since dates carved on turtles are
notoriously inaccurate—dates followed by the inscription B.C. are
sometimes found. This record, however, seems better substantiat
ed than most, and it can do little harm to accept it, bearing in mind
the circumstances. Despite this longevity, however, the maximum
size of an individual seems to be established after only a few years,
and very old specimens are often the same size or even smaller
than those whose clear-cut growth annuli establish their age as
less than ten years. When the adult size is reached, the carapace
bones fuse completely so that empty shells often show no signs of
the sutures between the individual bones.
Box turtles are omnivorous, many individuals showing a great
appetite for both live food (such as earthworms) and succulent
fruit. As with many turtles, there is a shift of diet from predomin
antly carnivorous to predominantly herbivorous with maturity.
Very few people ever eat box turtles, and those who do are appar
ently liable to be poisoned, since the flesh of a box turtle which has
fed upon certain varieties of fungi may be highly toxic.
Box turtles copulate shortly after waking from hibernation.
Courtship may be prolonged, involving biting of the shell and limbs
164
of the female. During actual intromission, the male is locked on to

the hind margin of the female’s shell by his hind feet, and he leans
over backwards so that any loss of grip will result in him falling on
his back. Nesting usually takes place in June and July, in late
afternoon or evening. Between two and seven eggs are laid (usual
ly four or five), which are about 1.25” long and 0.75” wide. There
are instances on record of retention of sperm by females for seve
ral seasons after copulation; Carr quotes an instance of a captive
box turtle laying five fertile eggs four years after any possible con
tact with a male. This phenomenon has also been reported for
diamondback turtles in captivity, and may possibly be of frequent
occurrence throughout the order.
In extreme north-east Florida and south-east Georgia, the
common box turtle intergrades with the Florida box turtle, Terra-
pene Carolina bauri. The shell shape shows no constant differences
from that of T.c. Carolina, though there is a tendency for the hind
margin of the carapace to flare more, and sometimes one encount
ers narrow, very high-shelled individuals which have no counter
parts among eastern box turtles. The coloration is much less vari
able than that of Carolina, the carapace having a striking pattern
The Florida box turtle, Terrapene Carolina bauri. Notice the great similarity of
pattern to Terrapene ornata. Photo by Ken Lucas, Steinhart Aquarium.
www.Ebook777.com
The two Mexican subspecies of the box turtle Terrapene Carolina: above, Ter-
rapene Carolina mexicana from the northeast; below, Terrapene Carolina
yucatana from Yucatan.
of narrow radiating yellow streaks on a dark brown or black back

ground. There is a continuous yellow streak along the midline of
the carapace. The plastron is usually almost unpigmented, but
there may be a few brown markings on some of the scutes. The
head is dark with a few yellow markings, including the constant
broken horizontal bar behind each eye. The average size is mar
ginally greater than that of the eastern box turtle, though the
record is only 6.75”. There are usually only three toes on each hind
foot.
The Florida box turtle is found throughout the peninsula, from
Alachua and Duval Counties through to the Keys; the highest pop
ulation densities are found in Dade County. The favored habitat
appears to be partially cleared mesic hammocks. Eggs are laid
from mid-April to June and number between one and nine (nor
mally about five).
During the Pleistocene, a box turtle known as Terrapene Caro
lina putnami, considerably larger than the living races, lived in a
wide area of the southern United States. It reached a carapace
length of over ten inches. It is thought that climatic fluctuations
and associated environmental changes during the Pleistocene led
to the differentiation of the early putnami stock into two modern
subspecies, the three-toed box turtle (Terrapene Carolina triun
guis) and the Gulf Coast box turtle (T.c. major). The three-toed
box turtle ranges from south-eastern Georgia to eastern Texas,
north to northern Missouri. The name is not a particularly useful
one, since some specimens have four toes, while the Florida box
turtle usually has three, as do some individuals of Carolina. This is
quite a small race, the usual carapace length being 4.5 inches, and
the record 6.5. The shell is narrow, elongated, and highly domed;
the posterior edge of the carapace is more flared than that of bauri
or Carolina, but less than that of major. The color is very variable;
however, in a typical phase the carapace is nearly uniform light
brown, with extensive red areas on the head and limbs, and white
lips. In another phase the shell may be covered with yellow spots
or with dark radiating lines, and the head with orange or yellow
spots. The plastron is often immaculate yellow. The habitat pre
ferences and diet appear to be similar to those of the eastern race.
The Gulf Coast box turtle, T.c. major, is a marsh-dwelling
form which exceeds in size all other American box turtles; adults
are often over seven inches long, sometimes over eight, thus ap
proaching the extinct putnami in size. The biggest ones are found
on St. Vincent Island, off the coast of the panhandle of Florida. The
Gulf Coast box turtle is found from the south-west corner of Geor
gia and the panhandle of Florida west along the coast to eastern
Texas. It is recognized by the strongly flaring rear marginals as
well as by the large adult size (most box turtles one finds are
167
www.Ebook777.com
adults, so the size difference has some value for identification).
The carapace may show patterns similar to those of bauri, Caro
lina, or triunguis, but in many individuals no pattern is present on
carapace or plastron, these being dark horn color or black. One
phase of this race has no counterpart among the other races; this
is an almost uniformly black animal, with a white face and snout.
The habits of this turtle appear to be fairly similar to those of Caro
lina, but the eggs are slightly larger and the males are said to en
gage in frequent fights during the mating season.
Around Thomasville, Georgia the box turtle population shows
influence of all four subspecies; no two individuals have the same
color pattern, and some beautifully marked specimens can be
found.
Two races of Carolina are found in Mexico: mexicana, in sou
thern Tamaulipas, northern Veracruz, and eastern San Luis Poto-
si, and yucatana in the northern part of the Yucatan Peninsula.
The first of these is closely related to triunguis, and the range of
the species was apparently continuous across Texas and Tamauli
pas in the late Pleistocene. Some specimens of mexicana have a
color pattern indistinguishable from that of triunguis, but there is
a tendency for the shell to be double vaulted, and occasional speci
mens have the white-blotched head of major. Some individuals
have a greenish cast to the shell and small black speckles on the
carapace scutes.
Terrapene Carolina yucatana has been collected at Piste and
Chichen Itza, Yucatan; Coba, Quintana Roo; and Dzibalchen,
Campeche. This subspecies is most similar to major, and its isola
tion from major presumably took place when sea levels rose at the
end of the last glacial period. The upper beak is deeply notched in
yucatana, and there are four toes on each hind foot (only three in
mexicana). The maximum shell length is about six inches. Many
specimens have the white-marked head shown by some southern
specimens of major. The carapace varies from olive-tan to dark
brown with paler blotches; the plastron may be yellow with brown
sutures or predominantly brown with irregular yellow areas. The
head is marked with black flecks, while the beak may have a
pinkish tinge.
When increasing aridity in south Texas and northern Mexico
forced a wide hiatus in the range of Terrapene Carolina, a small
population was isolated in an internal drainage area in central
Coahuila known as the basin of Cuatro Cienegas. This location is
essentially an oasis in the middle of a desert, and the box turtles,
findling life impossible in the arid surroundings of the basin, actu
ally entered the water and became secondarily—or rather tertiari-
ly—aquatic. This population is considered to warrant full specific
status under the name Terrapene coahuila. The Coahuila box tur-
168
tie has a small, narrow head, a uniformly brown shell with flared
marginals in large adults, and enlarged axillary scales. The throat
is mottled with grey and white. The plastron is unpigmented. This
turtle has not yet had time to perfect its adaptation to an aquatic
existence, and the shell is still somewhat elevated and the feet but
feebly webbed. The general effect is that of a mud turtle (Kinoster■
non), but of course the single plastral hinge immediately demon
strates its true affinities.
The natural history of the Coahuila box turtle was the subject
of a Master’s thesis by William S. Brown of Arizona State Univer
sity. Brown found that the preferred habitats of the species were
small, north-south trending, spring-fed marshes with shallow
water, mud bottoms, and dense aquatic vegetation. The marshes
are isolated and surrounded by desert grassland and scrub. The
turtles remain active throughout the year except for short periods
of extremely hot or cold weather. Mating occurs from September
to June, and eggs are laid from May to September. One to four
eggs are laid at a time, two or three being commonest, with the
average clutch containing 2.3 eggs. These clutches are distinctly
Comparison of the plastra of Terrapene Carolina bauri (left) and Terrapene or-
nata ornata (right). The carapaces of these two turtles are very similar.
169
www.Ebook777.com
■
smaller than those of U.S. Terrapene; however, about half of the

females may produce a second clutch later in the season, and
about a third of the females lay a third clutch.
The turtles forage in shallow water with the top of the shell out
of the water; the diet includes both insects and aquatic plants.
When picked up, they invariably retire into the shell and close up
tight until all disturbance is past; individuals of Terrapene Caro
lina and T. omata, on the other hand, may either close up or
struggle actively when picked up.
The population density of T. coahuila, within its restricted
habitat, is much higher than that of terrestrial box turtles and
more similar to that of many freshwater turtles; Brown found den
sities of 54 to 63 adult turtles per marsh acre.
The other section of Terrapene, the ornata group, includes two
species adapted for life in the comparatively arid south-central
and south-western United States and western Mexico. Numerous
rather minor morphological characters separate the two groups;
for example, turtles of the Carolina group usually have a postorbi
tal bar in the skull, a short interfemoral seam, a first central scute
that is concave along the midline, a carapace that has the highest
point toward the rear, and the bridge usually opposite the fifth
marginal. Turtles of the omata group, on the other hand, lack the
postorbital bar, have a first central scute that is straight or convex
along the midline (except in T. nelsoni), a carapace that has the
highest point toward the front and which is more flat-topped than
that of Carolina, and the bridge usually opposite the sixth margi
nal.
The eastern ornate box turtle, Terrapene omata omata, is
found throughout Texas except in the extreme east and west, north
to include all of Nebraska and the southern part of South Dakota,
and east to Illinois, where there is a disjunct colony on the northern
part of the border with Indiana. There is quite an extensive area of
overlap with the three-toed box turtle, but the species appear not to
hybridize. This turtle has a striking pattern of yellow lines radia
ting from the juvenile part of each of the large carapace scutes,
and this pattern is continued on the plastron, although in some
cases it would be more accurate to describe the plastral pattern as
consisting of dark brown lines on a yellow background. There are
yellow spots on the head, neck, and limbs. The shell is rather flat,
and there is no keel though there is a yellow line along the midline.
Typically this is a species of arid savannas and prairies, but occa
sionally specimens are found in swamps. Ornate box turtles
retreat at night and during hibernation into burrows; hibernation
begins with the first cold weather and may last from October to
May. A number of writers have remarked on the sudden appear
ance of literally thousands of ornate box turtles under certain wea
170
ther conditions, and population densities in some parts of the range

exceed those of any other box turtle. The eggs, three to five in
number, are laid in June and, as with the common box turtle, nest
ing usually takes place in the afternoon or evening. Like other box
turtles, this is an omnivorous form, but perhaps slightly more in
clined than others toward predation.
Between the Great Plains inhabited by the eastern ornate box
turtle and the western foothills of the Sierra Madre Occidental in
north-western Mexico there is a corridor through the Rocky Moun
tains in southern New Mexico and Arizona. This corridor became
alternately passable and impassable to many faunal elements dur
ing the Pleistocene, and it appears likely that the Salt Basin box
turtle, Terrapene omata luteola, which is still found in extreme
western Texas, northern Chihuahua and Sonora, and southern Ari
zona and New Mexico, is a survivor of the last exodus through the
corridor in the late Pleistocene. It is distinguished from the east
ern ornate box turtle by the absence of the plastral pattern; the
plastron is quite heavily pigmented but lacks radiating striae. This
subspecies may well be a relictual population that is on the way to
extinction because of a presently unfavorable climate; its popula
tion density is much lower than that of T. omata omata.
That population derived from omata which was isolated on the
western slopes of the Sierra Madre Occidental survives in appar
ently small and disjunct populations in Sonora, Sinaloa, and Naya-
rit, and has speciated and subspeciated into two forms, Terrapene
nelsoni nelsoni and T.n. klauberi. T.n. nelsoni is known from
Plastron of Terrapene
nelsoni klauberi from
Alamos, Sonora.
Specimen from the Col
umbus Zoo, Ohio.
171
www.Ebook777.com
nearly forty specimens from Pedro Pablo, Nayarlt. It has an elon
gate, rather narrow shell, a concave first central scute like that of
Carolina, and small coffee-colored spots all over the carapace and
plastron. Males have the medially turned inner toe, used for clasp
ing the female during copulation, that is also found in Terrapene
omata. From dissection of females it appears that the clutch size
is smaller, but the eggs themselves are larger, than those of north
ern box turtles; the average clutch numbers only 2.7 eggs, but they
are nearly two inches long and over an inch in width. Terrapene
ornata has an average clutch size of 4.7 eggs, but the average size
is 1.4 x 0.9 inches. Terrapene n. nelsoni appears to be restricted to
a very small area of oak-savannah habitat above 4000 feet altitude
near Pedro Pablo.
Terrapene nelsoni klauberi is found in the vicinity of Alamos
and Sierra de Batuc, Sonora, and near Terreros, Sinaloa; it is very
closely related to T.n. nelsoni, but has a rounder, less elongate
carapace, a nearly flat first central scute, and possibly a smoother,
less flared carapace in adult males. The spotted pattern is present
all over the shell and soft parts of the animal. It is possible that the
preferred habitat is less elevated and more forested than that of
nelsoni. No biological data are available for this race.
GENUS RHINOCLEMYS *
The genus Rhinoclemys includes nine terrestrial or semi-ter
restrial species whose collective range extends from southern Son
ora, Mexico, to Ecuador and Para, Brazil. They are larger than
box turtles and do not have the plastral hinge. They may be distin
guished from Clemmys not only by their mutually exclusive range
but by the structure of the carapace bones; the staggering of the
neurals with respect to the pleurals is so arranged that each neural
projects a little behind its associated pair of pleurals, with the re
sult that each neural bone has the shape of a hexagon with two
short sides adjacent to the posterior end; in Clemmys the neurals
project in front of their associated pleurals and the hexagonal
neurals have their short sides adjacent to .the front end.
The South American species of Rhinoclemys carry to a re
markable extreme the tendency among several groups of turtles
for southern representatives to lay fewer but bigger eggs than nor
thern representatives. All five of the South American species lay
eggs that are almost three inches long and V& inches wide, even
though the parent turtle is rarely more than eight inches long.
*Fretey, et al., 1977, Zool. Meded. Rijksm. Nat. Hist. Leiden,

52(6): 63-80, have reviewed the status of the names Rhinoclemmys,
Rhinoclemys, and Callopsis and decided that the correct name is
Rhinoclemmys. It has not been possible to always make this
change here.
Sometimes one finds dried Rhinoclemys shells containing eggs so

large that they cannot be extracted intact from any of the shell
openings. However, even though Rhinoclemys has no plastral
hinge, the plastron is not tightly co-ossified with the carapace at
the bridge, and during oviposition the bridge can give sufficiently
for the rear shell opening to allow the egg to pass through. As one
would expect, the eggs are so large that normally only one is laid
at a time.
This variation in egg number and size with latitude is quite
easy to rationalize. Turtles in far northern climes are under severe
environmental constraints throughout their entire annual cycle;
they have to hibernate for many months in winter, and there is
usually only one period of the year when eggs will be likely to get
sufficient warmth to hatch. Consequently only one, or very few,
clutches can be laid during the year, so these clutches must be as
large as possible to produce the numbers of young necessary to re
place the parental generation. The eggs will therefore be small
and as numerous as the body size of the female will allow. In the
tropics, however, the climate is such that eggs can be laid at any
time of the year and have a good chance of hatching. The tropical
turtle can then lay very big eggs, which produce correspondingly
big hatchlings which have a better chance of survival, because of
their size, than small hatchlings; and, although only one or two-
such eggs can be accomodated in the body at any one time, by fre
quent nestings in the course of the year it can produce as many
eggs as a northern turtle.
Rhinoclemys pulcherrima, sometimes known as the Mexican
red turtle or the Mexican wood turtle, is a most attractive species
with thin red lines on the face and extensive areas of red and black
vermiculations on the limbs, thighs, and tail as well as on the ven
tral parts of the marginal scutes and near the midline of the
plastron. It has a small head with finely serrated jaw edges. The
species as a whole ranges from extreme southern Sonora, Mexico,
to Costa Rica; it does not reach the Caribbean or Gulf drainages of
Mexico, but is known from the Caribbean side of Honduras. Four
subspecies are currently recognized, though variation is probably
more clinal than abrupt. The northernmost race was formerly call
ed pulcherrima pulcherrima, but Ernst has recently shown that the
name pulcherrima actually applies only to turtles from Guerrero
in southwestern Mexico. Guerrero turtles have an extensive
plastral figure, markedly striped bridge pattern, and a pleural pat
tern of only a single dark-bordered pale spot; the carapace is low.
R. p. pulcherrima is the least known of the subspecies and the one
with the apparently most restricted range.
R. p. rogerbarbouri is reported to range along the Mexican
Pacific coast from southern Sonora south as far as Colima,
173
www.Ebook777.com
though specimens are not currently available from extensive sec
tions of this range. The species appears to be moderately abundant
in Colima. Specimens from there are characterized by a relatively
low carapace, with a well-defined median keel, of an almost uni
formly brown color and with considerable surface sculpturing and
well-marked growth annuli. An adult carapace is about seven or
eight inches long and has a somewhat wedge-shaped dorsal aspect
with straight, posteriorly diverging sides whose slight revertion in
some specimens may possibly form channels for collecting rain
water for drinking, as happens with some species of tortoises. The
neural bones are typically hexagonal, the anterior ones with the
short sides behind and the posterior ones short-sided in front. The
plastron is yellow, with black bridges and a broad, black, some
what diffuse mid-plastral streak. The underside of each marginal
scute is marked with a red bar. In younger specimens both median
and lateral black markings are flanked by areas of red. The bridge
is poorly ossified posteriorly, a modification that may provide
some kineticism of the hind lobe to assist in oviposition. The limbs
are marked with mottling and longitudinal streaks of bright red
and black. Two red streaks cross from eye to eye above the nos
trils, and there is also a strong red line along each upper eyelid as
well as broken red lines on the sides of the head. The chin is bright
yellow, uniform below but with black mottling at the sides. The
eyes are a most attractive light blue, with a black bar through the
pupil. In the northern populations there are no red markings on the
head below the level of the eye.
Head of Rhino-
clemys areolata
from northern
Central America.
174
The populations from eastern Oaxaca, Chiapas, southern

Guatemala, El Salvador, and Honduras, currently called incisa,
are different in a number of ways. Most conspicuous is the much
deeper carapace of the adult (though hatchlings and young are
flat-shelled), and the oval rather than wedge-shaped appearance
of the shell from above. The carapace is again typically almost
uniform brown, though juveniles of Honduras specimens may
have dark-edged non-ocellar red streaks on the dorsal scutes. The
plastron is similar in color to that of the northern race. The red
striping of the face shows a number of differences, including the
frequent presence of a short anterior median streak and the pre
sence of pinkish-red streaks on both upper and lower jaw plates.
The chin is pale yellow and is mottled all over with black. The red
lines are in general a little thinner than those of rogerbarbouri,
and the V-shaped line extending from the snout to the ear region is
unbroken. There tend to be red markings on the sides of the head,
both above and below the level of the eyes. The snout in lateral
profile is almost vertical, while that of rogerbarbouri is notice
ably conical, with the nostrils distinctly anterior to the mouth.
Rhinoclemys pulcherrima manni, from Nicaragua and Costa
Rica, is an extremely decorative and distinctive turtle. It shares
with incisa the oval, domed shell, though again juveniles tend to be
rather flat. The carapace scutes are very strikingly marked with a
curving pattern of black lines and dots on a yellow background. In
young specimens some of the light carapace markings are bright
red. The plastron is yellow, with contrasting black bridges and
radial submarginal streaks and a variable black median area
which may be very narrow or may be broad with lateral projec
tions and isolated associated black spots. The chin is yellow with a
coarse pattern of black spots and circles. The head pattern in
cludes a continuous wavy V-shaped marking from the snout to the
back of the head, two red lines connecting the eyes and passing
above the nostrils, and other red bars on the jaws and sides of the
head. There may also be isolated dark-edged red spots on top of the
head. The surface of the carapace is smoother than that of the nor
thern race.
Rather little is known of the habits of the Mexican red turtle,
even though it is quite abundant in some areas. The habitat is al
most exclusively terrestrial, though during dry weather it may
move towards water and submerge itself briefly. It is good-
natured in captivity and feeds upon a wide variety of both animal
and vegetable matter. A typical clutch includes three brittle-shel
led eggs nearly two inches long.
Rhinoclemys areolata is a terrestrial species found in south
ern Veracruz, Tabasco, Campeche, the Yucatan Peninsula, north
ern Guatemala, and British Honduras (Belize). It is common on
175
www.Ebook777.com
Cozumel Island. The carapace is elevated, keeled, uniform grey-
brown in color, and has strongly sculptured scutes. Most individu
als are less than six inches long, but a few exceed eight inches. The
bridge is immaculate, but there is a large dark brown blotch cov
ering all but the extreme periphery of the plastron. The limbs and
tail are yellow with very small black spots. The upper part of the
head and snout are black, with a narrow bright red line on each
side extending back from the rear part of the eye. There may also
be small red markings above the eyes and on the back of the head.
Biological data for this species are lacking.
Rhinoclemys rubida is a completely terrestrial species found
in scattered localities in Pacific Mexico. Two subspecies are
recognized: rubida, which is known from many localities in Oax
aca, particularly on rocky hillsides near Tehuantepec, and also
narrowly entering the state of Chiapas; and perixantha, which is
known from tangled lowland scrub forest near Apatzingan,
Michoacan, and in various wooded coastal areas in southern
coastal Colima and also Jalisco and western Michoacan. At our
present state of knowledge, the gap between the most adjacent
populations of the two subspecies appears to be about four hundred
miles wide and includes the whole of the state of Guerrero.
This species is adult at about six inches and has a moderately
elevated, sculptured shell of a greyish-brown color, with a single
small yellow spot on each of the vertebral and costal scutes in well
marked specimens. The plastron is light yellow, with or without a
broad dark brown central blotch. The bridge is dark brown. The
head is relatively large and the upper jaw is non-serrated and
slightly hooked. The top of the head is characterized by a promi
nent horseshoe-shaped marking, with the apex forward, of a
creamy-white color. The pattern of the remainder of the top of the
head typically includes two elongate creamy-yellow spots located
close together and towards the rear; however, in many individu
als, possibly older ones especially, these spots may become fused
or completely irregular. There are two strong yellow bars above
the nostrils and other light lines on the sides of the head, the post
orbital streak being conspicuously broadened and sometimes
dark-centered. The limbs are mainly yellow, sometimes with ob
scure darker markings.
The two subspecies differ in shell shape, perixantha having a
flatter carapace with the marginal scutes projecting more out
ward than downward. In addition, the marginal scutes of perixan
tha contrast with the costals in being of a light tan color, the mar
ginals of rubida being an undifferentiated grey-brown. The gular
scutes are shorter, relative to the humerals, in perixantha than in
rubida. No data are available on the reproductive or dietary habits
of Rhinoclemys rubida.
176
Plastron of Rhinoclemys
funerea,an aquatic species
from eastern Central
America.
Rhinoclemys funerea Is the largest and most aquatic member

of the genus. It is found in the Caribbean lowlands of Costa Rica
and just enters adjacent parts of Nicaragua and Panama. It is re
ported from Bastimentos Island, Panama. This species is often
seen basking on logs in company with large Pseudemys. The cara
pace is low, streamlined, and can reach a length of at least thirteen
inches. In color the carapace is uniform black, though rapidly
growing individuals tend to be brown or greyish along the area
where the costal and marginal scutes come in contact. The plas-
tral scutes are also black, but each scute has a yellow margin. The
limbs, chin, throat, and sides of the head are yellow with black ver-
miculations; the top of the head is black with barely discernible
dark red-brown bands diverging backward behind each eye. The
feet are fully webbed. As with some other members of the genus,
when the head is fully retracted it is completely hidden behind two
flaps of skin stretched between the nuchal and gular scutes.
Sexual maturity in this species is reached at a length of about
eight inches. One to six eggs are laid at a time (average approxi
mately three). The eggs are hard-shelled and large, averaging
about 2.7” in length by 1.4” in width. The nesting season in
Panama extends from at least April to July, and one to four clutch
es are produced in a season. The species is herbivorous in nature
and feeds both in the water and on land.
177
www.Ebook777.com
Rhinoclemys annulata is a terrestrial species found in the
Pacific lowlands of Ecuador, north through the Choco of Colombia
at least as far as Barro Colorado Island, Panama. The type
locality of Chelopus gabbii, a name customarily synonymized with
R. annulata, was given as Sipurio, Costa Rica, by Cope in 1876; but
this town does not appear on recent maps of the country. This is a
dull-colored form, without bright markings anywhere on the body.
The carapace is uniformly mid- to dark brown, moderately eleva
ted, keeled, often flat-topped, and usually with some indications of
growth annuli. The plastron and bridges are dark brown to black
ish, with yellow bars connecting the fore and hind limb notches.
The limbs are short, stubby, and unwebbed and have an irregular
pattern of yellow and black. The jaws are white, but the top and
sides of the head are black with poorly defined yellow bars behind
the eye and on the sides of the neck. The hatchlings are much more
domed than those of other members of the genus and have a yellow
line along the midline. The carapace in adults measures between
six and eight inches, the record being 8.07 inches for a Colombian
specimen. Males have narrower shells than females, as well as
larger tails and concave plastra. It is possible, as with some other
members of the genus, that females may outnumber males; three
out of four Colombian adults examined by Medem were female, as
were six out of nine Panamanian adults examined by Mittermeier.
Despite its terrestrial habitat, this turtle drinks when the op
portunity presents itself and is sometimes found in rain puddles. It
relies entirely upon its cryptic coloration for protection and is not
known to dig even shallow burrows or to bite even when molested.
The natural diet on Barro Colorado Island includes ferns, shrubs,
and vines of a number of species; in captivity bananas are favor
ed. Trailing experiments by Mittermeier suggest that the normal
activity peaks extend from 7:00 to 10:00 AM and during and imme
diately after periods of rain. Fresh-caught specimens often carry
ticks (Amblyomma sp.), which in some cases excavate shallow
pockets in the shell of the turtle.
Mittermeier witnessed several copulatory attempts in this
species. The male would climb on to the female’s back, trying to
bite her head and insert his tail at the same time. The female
meanwhile would try to escape. After a few minutes of trying, the
male usually began to froth at the mouth. If the female is consider
ably larger than the male, head biting seems to serve only to scare
the female into her shell and reduce the frequency of escape at
tempts. Even if the male were to succeed in biting the head of a
larger female, it would do little good because, in such a position,
his tail could not reach hers. When the female is smaller than the
male, a successful head bite would appear to give the male more
leverage and thus facilitate copulation.
178
The single egg (rarely two) of annulata is about 2.75 inches

long and 1.4 inches wide; oviposition apparently takes place
throughout the year. The species is known as galapago to Spanish
speaking Panamanians, and paticua to the Cuna Indians of the San
Bias Islands. In parts of Colombia it is called montane.
Rhinoclemys nasuta is an aquatic species found in north-wes
tern Ecuador and the Choco of Colombia. This is another medium
sized species (record length 8.75 inches) with a rather low, dark
brown carapace. The plastron is yellowish with dark pigmentation
covering all except the anterior borders of each scute and a few
places along the midline; there are also pigmented areas on the
underside of the lateral marginals. The head and limbs too are
brown, with narrow white lines extending from the snout through
each eye to the neck and others running along the temporal mus
cles ending at the top of each eye. This species is found in large,
fast rivers, from which it only emerges to bask or to nest. In diet it
is principally herbivorous, eating aquatic vegetation, leaves, and
fruits. In captivity it usually refuses to eat and soon dies. As with
R. annulata a single huge egg is laid at a time, but oviposition pro
bably takes place throughout the year. The egg is not buried,
though sometimes a few leaves are pushed over it.
R. punctularia, in the broad sense, is a species group whose
range extends from eastern Panama south into western Colombia
and Ecuador, and east through northern Colombia, Venezuela,
Rhinoclemys nas
uta, an aquatic
species from
northwestern
South America.
179
www.Ebook777.com
Trinidad, Guyana, Surinam, French Guiana, and the Brazilian
Territory of Amapa and the State of Para. It is the only member of
the genus found east of the Andes.
Wermuth and Mertens (1961) consider five subspecies of punc-
tularia to be valid: punctularia in Trinidad, French Guiana, and
northern Brazil; diademata in Venezuela; lunata in Surinam;
melanostema in Colombia, Ecuador, and Panama; and nasuta in
Ecuador. However, not only does this arrangement not accord
with the facts (e.g. nasuta is also found in Colombia, while the spe
cies as a whole is also plentiful in Guyana), but it is also nonsensi
cal from the zoogeographic standpoint, for example in that
“lunata” is supposedly sandwiched between the Trinidad and the
French Guiana populations of “punctularia.”
I believe that nasuta and melanosterna, which are largely
sympatric and quite distinct in appearance, are not subspecifical-
ly related, and they are considered as separate species in this
book. Indeed, nasuta was considered a full species from the time of
its description by Boulenger in 1902 until 1954, when Mertens sub
sumed it within punctularia.
The form melanostema was described as a full species (of
Geoclemmys!) by Gray in 1861, but was relegated to varietal rank
within punctularia by Boulenger in 1889, who however only had two
preserved and discolored specimens available for his diagnosis.
Melanostema is distinguished by the very simple dorsal head pat
tern, which consists of two thin, sometimes interrupted, orange-
red dorsal lines which diverge posteriorly, and by the strikingly
light, almost white, irides, which contrast sharply with the pupils.
The neighboring Venezuelan form diademata has dark, non-con
trasting irides and a white or pale cream-colored dorsal head
marking which forms a pair of thick, wedge-shaped, posteriorly-
directed lateral streaks joined by a short, thick, curved crossbar.
Melanostema and diademata appear to be separate by the Sierra
de Perija that forms the northern part of the frontier between Col
ombia and Venezuela; melanostema has been recorded by Medem
from Santa Marta (Magdalena) and from La Girona near
Bucaramanga (Santander), Colombia, while diademata is
restricted to the western side of the state of Zulia, neighboring
parts of Tachira, and a small area of Norte de Santander, Colom
bia, in the Lake Maracaibo drainage basin. Therefore, in the
absence of any indication of intergradation between melanostema
and diademata, their close approach to each other’s ranges but ap
parent separation by the Sierra de Perija, and their morphological
distinctness, I believe it would be appropriate to recognize
melanostema and diademata as distinct species.
Rhinoclemys melanostema is known from the Colombian De
partments of Choco, Magdalena, and Santander, as well as from
180
Rhinoclemys punctularia from Trinidad showing the relatively large size of

the egg compared to the plastron length.
the Darien of Panama. It is a medium-sized turtle; males reach a

length of nine inches, females up to eleven inches. The shell is
moderately elevated and dark brown or black in color; the ground
color of the plastron is yellow, but an extensive dark brown or
black figure usually obscures all but the periphery. There are
rather complex dusky markings on the underside of the marginal
scutes. The skin of the throat, limbs, and tail is yellow with black
vermiculations. The top of the head is black, with two orange-red
streaks converging towards the snout, sometimes being broken
just above the eyes. The iris of the eye is pure white in juveniles,
very pale green in adults. In certain estuarine areas of Colombia,
the head stripes are pale green instead of orange. This species is
found in water or on land with comparable frequency. In the wild
state it lives almost entirely upon vegetable matter, but in captivi
ty shows a great appetite for meat and fish. As with other species
of this genus in Colombia, the clutch consists of a single elongate
egg of great size; the eggs are marginally larger even than those
of R. annulata. Medem found only four males in a sample of seven
teen specimens from Colombia.
181
www.Ebook777.com
As mentioned above, the form diademata should be considered
a full species. The type locality of diademata, given as Maracay,
Venezuela, is in error; the species is in fact restricted to the
western side of the Lake Maracaibo basin, bounded on the north by
the arid Guajira Peninsula, on the west by the Sierra de Perija, and
on the south by the Sierra de Merida. Little has been published on
this species, which is isolated from the nearest populaton of punc-
tularia by hundreds of miles of llanos. However, it has reproduced
freely in captivity in recent years at the Instituto Roberto Franco
in Villavicencio, Colombia, and appears to be quite abundant
within its restricted range. I picked up five dead juvenile
specimens on the highway from Maracaibo to Machiques in 1977.
The sex ratio appears to favor females strongly; I have examined a
series of ten live adults in the Maracaibo Zoo collection that only in
cluded one male, while the series in the La Salle Collection in
Bogota includes ten adult females, two adult males, and three
juveniles. The eggs of diademata measure 2.25 to 2.8 inches in
length and are about 1.3 inches in width; one to four are laid at a
time, at intervals of approximately two months. This is an om
nivorous species, feeding readily both on land and in the water.
The form lunata has recently been demonstrated by Fretey,
Hoogmoed, and Lescure (1977, Zool. Meded., Leiden, 52(6): 63-80)
to be invalid; it was based upon a specimen from Surinam with
unusually thick stripes on top of the head, joined anteriorly by a
rather thick crossbar. Fretey, et. al. located a total of four
specimens of the lunata phenotype and concluded that it was a rare
individual variation of punctularia, with no geographic or tax
onomic significance.
Rhinoclemys punctularia itself, then, is a much more
restricted species than normally thought, with no recognized
subspecies and a distribution that extends from Trinidad and ex
treme eastern Venezuela through Guyana, Surinam, and French
Guiana and into Amapa and Para, Brazil. A few specimens are
also known from Amazonas, Brazil (Lagoa Silva, Saracu, andLago
de Maximo), but it does not appear to extend very far into the
Amazon Basin. This species is characterized by the narrow,
posteriorly diverging red lines on top of the head; these are often
joined by a short crossbar in front, though this is not usually pre
sent in eastern specimens. I recently examined a group of about
thirty punctularia from Para, Brazil, all of which lacked the
crossbar. The plastron of punctularia is black, with yellow scute
borders; the carapace is uniformly black or very dark brown. The
head is small, with a rather pointed snout, and the neck and limbs
are vermiculated with a pattern of black and yellow. Not much is
known of its habits, but it is probably the most commonly seen tur
tle in much of the Guianas, where it lives in a wide variety of
182
ecosystems. Normally only one or two large eggs are laid, but I
once watched a small female, about six inches long, in the Guyana
Zoo lay three eggs, ranging in length from 1.9” to 2.1” and in width
from 1” to 1.1”. The eggs are not properly buried, but simply plac
ed in shallow depressions or crevices, possibly with a few dead
leaves pulled over them.
Mature females of all Rhinoclemys species in captivity appear
to be very liable to egg-binding, which often causes their deaths
within about a year. Young specimens, however, are very hardy,
and so presumably are adult males; but males, being much less
common than females, are hard to obtain._______________________
The overall distribution of the Emydidae is worth a little dis
cussion. The oldest known fossil attributable to this family is from
the Paleocene of North America, and eastern temperate North
America is one of the areas in which the family shows most diver
sity today. The other such area is tropical south-east Asia, and the
zoogeographer P.J. Darlington considers that the distribution of
living species suggests that the primary radiation of the family
was in this region. This seems reasonable, in vew of much higher
genus to species (and subspecies) ratio in south-east Asia than in
North America. The land zones between these two centers of
modern diversity, however, are very low in emydid species; only
two species are found in the western United States (though a few
more appear in western Mexico), and only three are found in the
entire Mediterranean area (in fact in the whole of Europe, Africa,
and the Near and Middle East); diversity begins as we reach the
India/Pakistan area and continues through to the Malay Archipe
lago. Emydids, in common with many other modern groups of ani
mals, failed to reach Australia and New Guinea. They are repre
sented by only two genera in the West Indies, Central America,
and all except extreme northern Mexico. The family has penetra
ted deep into South America, but has not prospered there; only
three species (two of them closely related) are found east of the
Andes, and none is found in the Amazon drainage system. Presum
ably they were displaced from, or prevented from entering, the
aquatic turtle niches in this area by competition from Podocnemis
and the chelid sidenecks, though in the northern hemisphere the
emydids prevailed and the sidenecks became extinct. In Africa
south of the Sahara, only one species is present (in West Africa);
presumably the long-established sidenecks (Pelomedusa and
Pelusios) already filled the aquatic turtle niches.
The three Mediterranean species are Emys orbicularis and
Mauremys leprosa and caspica.
GENUS E M Y S
Emys orbicularis, the European pond turtle, was once consid
ered to be congeneric with Blanding’s turtle, and both species have
183
www.Ebook777.com
slightly hinged plastra and a pattern of speckles all over the cara
pace. However, the neck, skull, and ribs of orbicularis show none
of the specializations found in blandingi, and the present separa
tion of the species into different genera is fully justified.
The European pond turtle is found in Spain, central France,
northern Italy, parts of southern Germany and Poland, west along
the Black Sea coast of Turkey to the Caspian lowlands in Iran and
part of Russia, and south to Morocco, Algeria, and Tunisia. The
adult length is five to six inches; the record is 7.5 inches. The shell
is rounder and flatter than that of Blanding’s turtle; the tail is very
long, especially in hatchlings. The pattern of the shell almost al
ways shows some light speckling, but in some specimens this may
be very slight, while in others the light color may actually predom
inate. Some individuals have the light speckles arranged into bro
ken radiating lines. The coloration of the plastron is very variable,
but the scute margins are usually black. Yellow spots or vermicu-
lations are present on the head, neck, and limbs of most individu
als. The European pond turtle is entirely carnivorous. It is some
times found a long way from water, but most individuals prefer to
swallow their food while submerged. The eggs are about one inch
long and 0.6 inch wide, and average ten per clutch. According to
Gadow, the nest site is prepared for digging by water from the
bladder, and the initial cavity is bored with the tail before the hind
feet are brought into action. The overall distribution of the Euro-
Juvenile European pond turtle, Emys orbicularis, from Russia. Photo by Dr.
Herbert R. Axelrod.
184
pean pond turtle appears to be decreasing, at least in Europe;

post-glacial remains have been found in Sweden, Denmark, Hol
land, and England. Adult individuals can live out of doors indefi
nitely in England, but they do not breed. Even in the natural popu
lations in some parts of central Europe, there may be total repro
ductive failure during cool summers.
GENUS MAUREMYS
The Spanish turtle, Mauremys leprosa, is found in Spain
(scarce in the north, common in the south), Portugal, Morocco,
Algeria, the Gambia, northern Mauritania, Libya, Tunisia, and
l’Adrar des Ifogha and the Air region of Mali. There are also rec
ords from southern Dahomey, which would suggest that the spe
cies has a very wide distribution in tropical West Africa; on the
other hand, Dahomey records are rejected as unreliable by Love-
ridge and Williams. Extremely high population densities are
reached in some parts of the range. This species is easily distin
guished from Emys by the unhinged plastron and the seam be
tween the femorals being longer than that between the anals. The
carapace is depressed and the nuchal scute is wider than long.
Young specimens are almost circular and have three low keels.
The coloration is somewhat variable, but young specimens are of
ten a rich brown color with an orange figure on each costal scute.
The ground color of the plastron and under-marginals is pale yel
low, but an extensive dark brown or black area usually obscures
all but the periphery of the plastron; the underside of the margi
nals also shows extensive black pigmentation, and there is a black
streak across each bridge. The soft parts are olive-grey, with pale
yellow stripes along the neck and limbs and a greyish-orange cir
cular spot present behind each eye. Large adults (which may mea
sure six or seven inches in length) lose the juvenile keels and color
ation and become a uniform dull greyish color. They also develop
somewhat enlarged heads. When picked up, freshly caught Span
ish turtles emit a fishy-smelling liquid from their inguinal glands.
Mauremys leprosa is aquatic and carnivorous. Although
sometimes found in clean water, they often occur in filthy mud-
holes, and under these circumstances are most subject both to in
festation with parasites and epizootic algae and to various ero-
sional diseases, from which the specific name ‘leprosa’ is derived.
Eggs, usually numbering from six to eight, are laid in early May,
according to Louis Lortet, and hatch only 25 days later. The eggs
are larger than those of Emys, measuring about 1.4 inches x 0.8
inches. In captivity these turtles are hardy, but males have a ten
dency to bite the skin around the neck of other turtles, usually of
females of their own species, and the sores that develop may be
difficult to heal. When mating, the male turtle holds on to the cara-
185
www.Ebook777.com
Mauremys leprosa, an aquatic species of northern Africa and Iberia. Photo by
L.E. Perkins.
pace of the female with all four feet, holding on so tight that the fe
male may he picked up and even inverted without the male falling
off, and while in this position the male still makes rather vicious
bites at the female’s head and neck.
The Caspian turtle, Mauremys caspica, is quite closely related
to Mauremys leprosa. Two races of this species exist: M.c. cas
pica in western Iran, extending north into Armenia and Dagestan,
south at least to Shiraz, and in north-east Arabia at least to Bah
rain ; eastward through Iraq into eastern Turkey, where it is re
placed by M.c. rivulata, whose range includes southern Yugo
slavia, Albania, Crete, Greece, some of the Aegean Islands (Corfu,
Milos, Kythnos, Kos, Salonika, Chios), southern Bulgaria, Turkey,
Syria, Israel, and Cyprus. The typical race reaches a length of
about eight inches (record 9)4 inches) and has a depressed cara
pace and a small head even in large adults. The carapace, which is
slightly tapered and pinched upward in the tail region, is red-
brown vermiculated with olive-green; its surface is smooth in
186
adults but wrinkled in young specimens. The plastron is notched

behind and is predominantly black with yellow present along the
seams. The bridge is basically yellow, with a row of black spots,
two to each scute, along the lower marginals, and with black
streaks mainly confined to the seams. The head is basically olive-
green, unmarked above except in the juvenile, when it bears an
asymmetrical pattern of light lines. A yellow streak runs from the
nostril to the eye and is continued behind the eye and along the
neck. A broader streak from the nostril ends below the eye. Ano
ther pale line extends downward from the nostril to the cusp of the
jaw and is continued along the jaw; there is a faint line just above
this one. The lower jaw is marbled with white markings which
form longitudinal lines along the neck posteriorly; broad pale yel
low streaks are present in the axillary and shoulder regions. The
forelimbs are dark olive-green with yellow stripes, two particu-
Mauremys caspica caspica from the Near East and Turkey. Photo by G. Mar
cuse.
larly broad ones, confluent for most of their length, extending to

the inside two digits. The color pattern is maintained even in large
adults. This is an abundant species, and although much of its habi
tat is very arid country, large basking aggregations may be seen
in almost any body of water within the range. Males reach the
same size as females, but can be recognized by the usual charac
ters of the slightly concave plastron and more distally located
vent. The eggs, probably usually four or five in number, measure
about 1.6” x 1.2”.
Mauremys caspica rivulata differs from the typical race in
having a completely black plastron, almost entirely black bridges,
and the head dark brown on top, with a dark streak passing from
the eye to the tympanum, and the remainder of the head pale
187
www.Ebook777.com
brown with pale linear markings; the head coloration of the juve
nile is similar to that of M.c. caspica. This turtle is reported to live
in marshes, ponds, lakes, and slow rivers. The female lays four or
five eggs about the end of July. It is possible that the Crete popula
tion constitutes a different subspecies (cretica) distinguished by
the lighter pigmentation and brighter iris.
Mauremys mutica, an eastern relative of the Caspian turtle, is
found in northern Indo-China and in mainland China as far as
Kiangsu and Anhwei Provinces, as well as on Hainan and Taiwan.
This species reaches a length similar to that of mature M. caspica,
but the carapace is conspicuously deeper and has a strong median
keel with weaker lateral keels. The carapace is brown with black
seams; the plastron is basically orange-yellow, but a large black
blotch almost obscures each individual scute (though the gulars
are sometimes unpigmented); two more black smudges are pre
sent on each bridge, and smaller markings are on the undersides
of the marginals. The head is olive above, yellow below, and with a
broad yellow band extending backward from the eye to the tym
panum, the upper margin of which is dark. Mauremys nigricans is
a synonym of mutica, based on two specimens with black pigmen
tation on the chin and throat. Mauremys mutica shows great vari
ability in the shape of the neural bones; in some specimens the
hexagonal neurals have their short sides adjacent to the front mar-
The Asian Mauremys mutica, once commonly called Clemmys nigricans.

Photo by G. Marcuse.
188
Carapace and plastron of Mauremys japonica, a nearly all-brown species.
gin, in others adjacent to the hind margin. Very little is known of

the biology of this species beyond the fact that it appears to be an
aquatic species inhabiting low altitudes. It is very hardy in cap
tivity and makes an excellent pet.
Further east still, Mauremys japonica is found in the southern
parts of Japan, north to around the level of Tokyo. This species has
an adult length of about five or six inches. The carapace is brown,
very depressed, and has a median keel. Each scute shows both
growth annuli and radiating striae. The plastron is black. The tail
is long—half the length of the carapace in adults, about equal to the
carapace length in hatchlings. The skin is olive-brown or blackish,
with a pale streak along the outer border of the limbs and a broad
pale band along the upper surface of the tail. The top of the head is
uniform brown, and the general dorsal effect is similar to that of a
narrow-shelled version of Cyclemys dentata.
The turtles included in the genus Mauremys were until recent
ly considered to belong to the American genus Clemmys. How
ever, there are numerous skull features that separate the two
groups, and their similarities are probably the result of a similar
way of life rather than of close relationship. On the other hand, if
similarities in courtship pattern are taken to indicate phylogenetic
relationship, it must be remembered that males of both Clemmys
189
www.Ebook777.com
insculpta and Mauremys caspica are exceedingly violent lovers,
biting the necks of their females with such viciousness that the
wounds inflicted, under captive conditions, are often fatal.
GENUS SACALIA
In addition to the species already described, one other species
which was formerly included within the genus Clemmys, but
which is now considered distinct, is Sacalia bealei. This species is
about five inches long when adult, and occurs in southern China
(from Fukien to Hainan Island) and northern Indo-China. The
carapace is moderately depressed and with a median keel toward
the rear. The plastral buttresses are very weak. The axillary
scutes are small and the inguinals vestigial or absent. The shell is
speckled or vermiculated with black on a yellowish-brown back
ground. The plastron, soft parts, and eyes of living specimens from
Futsing Hsien are pink, according to Clifford Pope. The most no
ticeable feature of the species is the presence of two or four
striking ocelli on the temporal region of the head. The hind pair is
always present, but the front ones may be obscured by black ver-
miculation on the greyish-olive ground color of the head. The front
faces of the forelimbs are yellow, while the lateral aspects are
brown. The species Sacalia quadriocellata, from Fukien, Kwang-
tung, and Hainan Island, is considered by Pope to be distinct from
bealei, though in their checklist Wermuth and Mertens synony-
Plastron of Sacalia bealei of China and northern Southeast Asia. Photo by Dr.
Robert Mertens.
190
Striking head pattern of a turtle agreeing with the description of Sacalia

quadriocellata.
mize the two forms. This form is reported always to have four
black-centered white ocelli in the temporal region, the hind ones
being the larger, and to lack the contrasting yellow and brown
color of the forelimbs found in bealei. The carapace is dark brown
with dark radiating lines more or less evident on each scute. The
plastron is yellow, profusely covered with small, dark spots, many
of them elongate. There are yellow stripes on the neck. S. quadri
ocellata is reported to be an inhabitant of mountain streams, prob
ably preferring wooded regions. Only two eggs are laid at a time.
GENUS ANNAMEMYS
A turtle considered somewhat related to Mauremys, but dif
fering from that genus in the much greater strength of its plastral
buttresses, is Annamemys annamensis; the somewhat redundant-
sounding scientific name of this species is the result of the fact that
it was first described under the name Cyclemys annamensis, and
later and independently by the name Annamemys merkleni. Since
191
www.Ebook777.com
it does not properly belong to the genus Cyclemys, the later
generic name Annamemys must be used, but the earlier specific
name remains valid. This species has only been recorded from
Phuc-Son and Fai-Fo, Vietnam. It has a rather low, uniform grey-
brown carapace, tricarinate in young specimens, that reaches a
length of about six inches. The ground color of the plastron is yel
low or orange-yellow, with a large black blotch almost obscuring
each scute, and further black blotches along each bridge and on
the lower marginals. The upper part of the head is dark brown; a
pale yellow stripe begins at each nostril and passes through each
eye and along the neck; this line reaches its widest point where the
skin of the head meets the soft skin of the neck. Other pale yellow
lines pass below each eye; the chin and throat are creamy-white.
The toes are completely webbed. There is a notch at the mid-point
of the upper jaw. The plastral buttresses are well-developed,
nearly reaching the neural bones. The front feet have five claws
each, the hind feet four. No field observations on this species have
been recorded.
Several genera of Asiatic emydids have even stronger plastral
buttresses than Annamemys; these buttresses enclose large
192
chambers at each side of the shell, which in life are occupied by the
lungs. Turtles with these lateral lung chambers are typically large
diving forms which remain submerged for considerable periods of
time and thus require voluminous lungs. Four of the five genera
(Hardella, Batagur, Callagur, Orlitia) each include a single very
large species, while the fifth (Kachuga) comprises seven species,
some of which are very large and others quite small.
GENUS HARDELLA
Hardella thurji, the Brahminy river turtle, is found in the Gan
ges, Brahmaputra, and Indus River systems, ranging from Sind
eastward to Sylhet; in Pakistan it ascends the Indus at least to
Sukkur. Large numbers are sold for food in Calcutta. It is a large
species, females reaching a carapace length of at least 21 inches,
though males are a great deal smaller (reportedly less than seven
inches long—if this is true, this species shows the most extreme
sexual dimorphism among the Chelonia). Young specimens have a
rather deep carapace with a slightly wavy median keel. In color
the carapace is very dark brown or black, with a yellow margin in
all but very large individuals and a yellow band along the junction
Hardella thurji, a common species of India and Pakistan. Photo by G. Mar

cuse.
193
www.Ebook777.com
of the costals with the marginals in rapidly growing specimens.
The plastron is pale yellow with two large black blotches on each
bridge, another black area on the midline of the plastron near the
front, large blotches on each abdominal scute, with smaller exten
sions on the posterior parts of the pectorals, and more black pig
mentation around the interfemoral and interanal seams. The head
is large, black in color, and the snout projects somewhat beyond
the upper jaw. A strong yellow stripe on each side extends from the
nostril, passes above each eye, and then curves sharply downward
along the side of the head and neck. This stripe is sometimes bro
ken above the eye. Another elongate yellow blotch is situated just
below the nostrils, the lower jaw is pale yellow, and there is a
short, diagonal yellow bar just below each eye. The limbs are
black and lack a striped pattern. In young specimens the central
scutes are wider than long, and in adults they are only a little nar
rower. The tail is always short. The digits are completely webbed.
The Brahminy river turtle is highly aquatic, rarely being seen
basking, but prefers still ponds, canals, and sluggish backwaters
to fast-flowing rivers. In diet this species is herbivorous, and in
captivity fruit and vegetables of many kinds are eaten. It shows no
inclination to bite when handled.
GENUS BATAGUR
Batagur baska, the common batagur or tuntong, is distin
guished from related genera by the presence of a denticulated
middle ridge in both upper and lower jaws and by the presence of
only four claws on each front foot. This is a very large species
reaching a maximum length of about 23 inches. It is found in
Sumatra, the Malay Peninsula, and north through Thailand to the
Irrawaddy River in Burma; it is frequently found in brackish, or
even salt, water. The general appearance of the batagur is very
similar to that of the Central American river turtle, Dermatemys',
the shell is uniform brown, smooth and heavy, and with very little
indication of scute boundaries in large specimens. The head is also
brown, with a strongly projecting snout. The plastron is orange-
yellow and of course lacks the inframarginals of Dermatemys. In
the breeding season, the male assumes a much brighter colora
tion, the nostrils becoming pale blue but the rest of the head in
tense black, passing into a rich crimson at the neck; this crimson
color extends over the whole of the front limbs. The batagur lives
in large rivers and nests colonially on sandbanks. In the Irrawad
dy, nesting takes place from the beginning of January to early
March; the eggs measure about three inches in length and slightly
more than half that in breadth. Each female usually lays three
clutches during a season, at about fortnightly intervals, each
clutch numbering from ten to thirty eggs; the total season’s pro
duction per female averages 50 to 60. The incubation period is
194
about seventy days. The eggs are eagerly collected by the local
human population, and it is probable that batagurs are nowadays
much less abundant than formerly. In the Malay Peninsula, the
batagur is found right down to the southern tip, but the most im
portant nesting beaches are in the Perak River near the west
coast. A few batagurs also nest on the leatherback nesting beach in
Trengganu. Under local law, one-third of the eggs collected in
Perak by licencees must be allowed to hatch, and the young reared
until they are large enough to be safely released, the government
suppying the funds for feeding the young turtles. However, the tur
tles are decreasing rapidly, possibly because of poor enforcement
of the conservation regulations.
The information given above on the batagur was taken largely
from two very old publications—“Anatomical and Zoological
Results of the Yunnan Expeditions. Reptilia and Amphibia” by
John Anderson (1879) and “Reports on Inland and Sea Fisheries in
the Thongwa, Myaungmya, and Bassein Districts and Turtle-
Banks of the Irrawaddy Division” by F.D. Maxwell (1911). Little
research effort seems to have been directed at the species since
then, even though batagur eggs are a valuable item of commerce
In Malaysia. During the past year, however, Edward Moll of
Eastern Illinois University has undertaken a survey of the Perak
River population of batagurs on behalf of the World Wildlife Fund.
His findings have not been published as of the time of writing, but
Moll writes me that he can already see many errors and question
able conclusions in the older literature. For example, in Malaysia
breeding male batagurs acquire an intensely black coloration on
the head, neck, and limbs, and the eyes become bright white, but
the red coloration mentioned by Anderson was not observed. How
ever, Moll did perceive some geographic variation in color pattern
between the western and eastern coast populations in Malaysia.
Moll summarized his batagur findings in remarkable poetic
language in the World Wildlife Fund Special Report for Summer
1976:
Pekan Baharu, Malaysia: The nesting migration is

easily the most spectacular event in the life history of the
Malaysian river terrapin, Batagur. A 50 to 60 mile journey
that begins between November and early January takes the
turtles up the Perak River to nest on an extensive area of
sand banks and islands.
With the eggs deposited, the terrapin scrapes sand over
them and then compacts the sand by repeatedly dropping
her pound hulk upon it. The resultant sound of “tun tonk”
has led to the terrapin’s Malay name of “tuntong.” As
twenty to thirty terrapins drum the sand, the sound carries
195
www.Ebook777.com
up and doum the Perak (River) announcing, as it has for
eons, the annual return of the Batagur nesting spectacular.
The sound is weaker now than in earlier years, for now the
drummers number in the hundreds instead of the thousands.
Though the sound has diminished, it has not vanished, and
there is hope that it will again resound across the Perak at
former intensities.
GENUS CALLAGUR
Callagur bomeoensis is found in the Malay Peninsula, Borneo,
and Sumatra; its local name in Malaya is biuku. It may be the lar
gest of all emydids; according to Nelly de Rooij, it reaches a cara
pace length of about thirty inches. It differs from Batagur in hav
ing five claws on each forelimb, from Orlitia in having the top of
the head covered with undivided skin instead of small scales, and
from Kachuga by the elongated neural bones and narrow jaw sur
faces. Young specimens are grey all over, both shell and skin, the
grey color being lightest on the plastron; there are no spots or
stripes of any kind. However, with age the carapace becomes
grey-brown with three black longitudinal bands, while the under
side loses all grey pigmentation except under the marginal scutes.
The head is rather small and the snout projects beyond the upper
Very small specimen of Callagur borneoensis, a species seldom available in

the United States.
jaw—more so in adults than in young. The tail is short and the

digits are completely webbed. Young specimens have rather flat
tened shells with strong median keels; the marginal bones, parti
cularly toward the rear, are very wide, adding considerably to the
width of the shell. The toes appear very long and delicate. Callagur
bomeoensis is very closely related to Kachuga and might be con
sidered as a southeastern representative of that genus. Its habits
have not been studied, but Mertens reports that in Sumatra it
shows a preference for brackish water. This species has been ques
tionably reported from the Philippines.
GENUS ORLITIA
Orlitia bomeensis is another huge species found in the Malay
Peninsula, Borneo, and Sumatra. The maximum length is around
28 inches. Young specimens have a rather humped, uniform black
carapace and black mottlings on the head. The underside is pale
buff, with a dark rim around the under-marginals in young speci-
Carapace of Orlitia borneensis. Specimen in Senckenberg Museum.
mens. A white line extends back and down from the comer of the
mouth. Adult specimens have a rather low, narrow, smooth shell.
The head is large (distinctly larger than in Callagur), short, broad,
and the upper jaw is hooked. The skin of the back of the head is
197
www.Ebook777.com
covered with small scales. The scales on the faces of the forelimbs
are broadened and band-like. The toes are completely webbed, in
keeping with the thoroughly aquatic habitat. The skin between the
eye and the tympanum and on the underside of the tail is granular
(covered with small scales in the larger Kachuga species).
GENUS KACHUGA
Kachuga includes seven species, some of which are extremely
large in size, others quite small. The most familiar is Kachuga
tecta, the Indian roofed turtle, which is found in the Ganges, Brah
maputra, and Indus River systems of northern India; in Pakistan
it appears to be restricted to the Indus Valley of middle Sind within
a 50 mile radius of Manchar Lake. This is a small species, being
adult at about seven inches (males less); the record length is nine
inches. The general shape is strongly reminiscent of that of the
American sawback turtles (Graptemys)\ the carapace is very
elevated, with flat, tent-like sides and a strong, tuberculate
median keel which reaches its highest point at the rear of the third
central. As in most members of the genus, the central scute row is
strongly constricted at the junction of the third and fourth cen
trals, and the fourth central is strikingly elongate. The carapace is
brown, with a red median stripe, and the plastron is pinkish-yellow
with two or three black blotches on each scute which form a sym
metrical pattern. There are other black streaks and spots on the
bridge and under the marginal scutes. The rim of the shell is
yellow or pink. A red marking behind each eye is of similar shape
to the yellow postocular marking in the yellow-bellied turtle (Pseu■
demys scripta scripta). The neck is decorated with many fine
longitudinal lines, and the limbs are spotted. This is a quiet water
species and is apparently vegetarian in diet. According to Sher
man Minton, it is less alert and a poorer swimmer than K. smithi.
Kachuga tentoria is often considered a subspecies of K. tecta,
but no intermediates have been found even though the two species
occur together in Bangladesh. K. tentoria is found in peninsular
India, being known from the Mahanadi, Godavari, and Kistna
River systems. The adult size is similar to that of tecta; it is distin
guished from that species by the short second central scute, the
unpatterned limbs, the fewer and less distinct neck stripes, and the
absence of a large crescent marking behind the eye (a small pink
spot only is present). The plastron is pinkish buff, with coalescing
black spots along the submarginals, and usually only a single ir
regular black blotch on each of the plastral scutes. There is a longi
tudinal pink stripe along the middle of the carapace. Each bridge
is decorated with a broken longitudinal streak, and a small black
spot is located on each axillary and inguinal scute. A newly de
scribed subspecies (K.t. circumdata), from the Calcutta region, is
198
distinguished from the typical race (K.t. tentoria) by the presence

of a pink band along the junction of the costal and marginal scutes.
Kachuga smithi is found in the Indus and the western Ganges
system. The overall appearance is that of a low-shelled, pallid
skinned Kachuga tentoria. The carapace is brown with a dark
brown or black median keel, only feebly tuberculate. The first two
centrals are very short, the third and fourth strikingly elongate
and only very narrowly in contact. The plastron is completely
black except for narrow white margins to each of the scutes. The
skin is pale buff or grey, darker on top of the head, and there is a
prominent pink or red spot behind each eye. The neck is longitudi
nally striped. The snout projects well beyond the upper jaw, the
nostrils being perforated below the tip. The jaws are serrated, with
a large terminal cusp on the lower jaw. The digits are fully
webbed, and the scales on the limbs are transversely enlarged.
This is a common species in Bangladesh south of Jhelum, and
basking aggregations are often seen. The maximum length is
about nine inches. A female recorded by Sherman Minton laid
seven eggs, each 1.8 long by 0.9 inch wide. This species is a rapid
swimmer and is more carnivorous in diet than K. tecta.
Kachuga dhongoka is found in the eastern Ganges system as
far west as Allahabad and north to Nepal; it is also known from the
Brahmaputra. Females reach a maximum length of about sixteen
Kachuga dhongoka, a seldom-seen species of the Indian region.
199
www.Ebook777.com
inches, males about ten inches. The shell is brown and rather low,
but the median keel is well-defined, tuberculate, and marked by a
black line. The plastron is narrow, yellow in color, with a large
reddish-brown patch on each scute in the young. The soft parts are
brownish or yellow, and a white or light yellow line runs from each
nostril, through the eye, and ends at the back of the head behind
the mouth. The eggs number around 30 to 35 per clutch and are
about 2.2 inches long by 1.3 inches wide.
Kachuga kachuga is found in the Ganges system in Bangla
desh, Nepal, Bihar and Uttar Pradesh, and possibly also in the
Kistna and Godavari Rivers. Females reach a length of 15 or 16
inches, males less. The shell is low, and the median keel is most
prominent on the second and third central scutes; a tubercle at the
rear of the third central is present in young specimens but disap
pears with age. The third and fourth centrals have a fairly wide
line of contact. The carapace is olivaceous or brownish in color,
the plastron yellowish. Males in breeding condition have seven red
or red-brown longitudinal stripes along the neck, a pair of oblong
red or yellow spots on the throat, and the head marked with a bril
liant red on the top and with bluish on the sides. In the Ganges, egg
laying is reported to take place in March.
Kachuga trivittata is a very large species from Burma, known
from the Salween River and the Irrawaddy as far north as Bhamo.
Females reach a carapace length of over 23 inches, males about 18
inches. This species is closely related to K. kachuga, but the ridge
on the upper jaw surfaces is closer to the inner than the outer mar
gin (closer to the outer in kachuga). The male has a pale olive-
green carapace with three conspicuous longitudinal stripes which
may be united at their extremities. The plastron is pale orange-
yellow. The female is dark brownish, both above and below. The
head and neck are olivaceous, with a large black lozenge-shaped
patch or streak on top of the head. The limbs are yellowish. The
eggs are said to average 25 in number, about 2.9 inches by 1.6
inches in size, and are deposited in sandbanks above tidal limits in
December and January.
Kachuga sylhetensis is a small, rare species from the Garo,
Khasi, and Naga Hills in Assam. The shell reaches a length of
about seven inches. The carapace is strikingly elevated, and the
strong median keel is produced into a very strong backwardly pro
jecting spike at the posterior margin of the third central. The first
two centrals are very short, the third and fourth long and with a
very narrow line of contact. There are 24 marginal scutes (not
counting the paired supracaudals), two more than in the other spe
cies of the genus. The carapace is olive-brown, the median keel
paler; the plastron is yellow, each shield with a large dark spot.
There is a yellow wavy transverse stripe across the back of the
200
head and another along the lower jaw, and there are longitudinal
light streaks on the neck.
GENUS GEOEMYDA
For many years the genus Geoemyda was held up as an exam
ple of an emydid genus whose species were found in both the New
and the Old Worlds, and the sixteen or so species made it one of the
largest genera of the family. However, a recent revision of the
group by Samuel McDowell shows that the American species con
stitute a separate genus (Rhinoclemys), and that even the Asiatic
species are so diverse that they in fact represent three genera
(Geoemyda, Heosemys, and Melanochelys), differentiated not
only by superficial differences but by quite basic skull features.
The only features that all of the American and Asiatic species of
Geoemyda (in the old sense) share are the hexagonal neural
bones, which have the two short sides adjacent to the posterior
margin, and the tendency to abandon a strictly aquatic habitat.
Geoemyda spengleri is now the only species left in this genus.
It is found in the provinces of Kwang Si and adjacent Kwangtung in
China, in Viet Nam, in certain of the Indonesian Islands, and on
Okinawa. The Okinawa population is designated as a separate sub
species (japonica), distinguished by the presence of axillary
scutes and sometimes inguinals as well (both are absent in the typ
ical race). This is a small species, about five inches in carapace
length, and the shell is depressed, tricarinate, and serrated both in
front and behind. The upper jaw is hooked, and the skin on the back
of the head is smooth. The overall dorsal coloration is brown, more
or less variegated with blackish or yellowish; the underside is
black, with yellow on the bridge, the serrated ends of the margi
nals, and the margin of the plastron. The head and limbs are
brownish, with a yellow streak from behind the eye to the neck,
and yellow spots on the snout, tympanic region, and throat. Al
though basically a terrestrial species, the digits are partially web
bed. According to Clifford Pope, its abundance in Yaoshan may be
taken as evidence that it likes wild, wooded, mountainous country.
GENUS HEOSEMYS
The genus Heosemys includes five species, all of which lack
side keels but have prominent median keels and lack a bony tem
poral arch in the skull.
Heosemys grandis is a very large species; it is found in Burma
as far north as Pegu; Thailand; Cambodia; and the Malay Penin
sula as far south as Province Wellsley. The carapace is very broad
in young specimens, more elongate in adults, and reaches a length
of up to 17 inches. The shell is moderately elevated, brown in color,
usually with well-defined concentric annuli on each scute, and with
201
www.Ebook777.com
a pale brown median keel. The plastron and bridges are pale buff
with a very striking pattern of radiating dark brown lines on each
scute; either the black or the buff color may predominate. The
head, neck, and forelimbs are quite massive. The head is brown,
with small buff or pinkish spots and vermiculations on the sides;
the upper jaw is strongly bicuspid at the front. The forelimbs are
brown and covered with scales, some of which are enlarged. The
toes are webbed. Young and half-grown specimens have a large
fontanelle or ‘soft spot’ in the middle of the plastron. This species
appears almost equally at home in water or on land, eats almost
anything edible, and grows very rapidly under good conditions.
Heosemys spinosa is a bizarre species from Tenasserim, pen
insular Thailand, the Malay Peninsula, Sumatra, Borneo, and the
Natuna Islands. Hatchlings are about 2.5 inches long and have a
uniformly brown carapace which is rather flat and nearly as wide
as long. The median keel is extremely strong, paler in color than
the rest of the shell, and nearly one-third the width of the central
scutes. This keel forms two small spines at the rear of the second,
third, and fourth central, and a single spine at the rear of the fifth.
There is also a small but sharp spine near the middle of the hind
margin of each costal scute, and the exterior margin of each mar
ginal scute is drawn out into a prominent spine, so that the total
effect is that of a cogwheel. There is also a row of very small spines
between the anteriormost large marginal spines, on the first pair
of marginal scutes, and a secondary spine is present on some or all
of the last four marginals on each side (but not on the supracau-
dals). The plastron is pale uniform buff. The head is bluntly point
ed and the upper jaw is bicuspid, as in H. grandis. The basic color
of the soft parts is chocolate brown, but the sides of the head and
many of the scales on the limbs and tail are marked with pink. The
most obvious changes with maturity are a gradual deepening and
slight narrowing of the carapace, up-turning of the marginal
spines, and formation of a most attractive pattern of fine rich
brown radiating striae on each of the plastral scutes and oh the
underside of the marginals. The spiny turtle reaches a length of
eight or nine inches. In Penang and Perak, Malaya, it is found in
jungle hill-streams at altitudes of between 2000 and 3000 feet, but in
Singapore it is found on Bukit Timah at less than 500 feet. It is said
to be aquatic, but since the toes are virtually without webs it is
poor at swimming. It is herbivorous in the wild state, but in cap
tivity will eat either vegetable or animal matter. The function of
the spines is not known; possibly they lessen the likelihood of the
animal being swallowed by snakes.
Heosemys silvatica is a rare and poorly-known species from
the dense forests of Cochin, in southern India. The type specimen
(a male) had a carapace length of 4.7 inches. The carapace is de-
202
The shape, spiny marginals, and radiated plastral pattern are very distinctive
in Heosemys grandis. This species is often kept in captivity. Photos by M F
Roberts.
pressed and traces of lateral keels are present. THe posterior mar
gin of the carapace is feebly reverted, but is not serrated. The head
is rather large and the upper jaw hooked. The skin on the back of
the head is divided into moderately large scales. The forelimbs
have enlarged squarish or pointed horny scales on the front face;
the hind limbs are club-shaped, and the digits are virtually unweb
bed. The carapace is dark bronze color and the plastron yellowish
with two dark blotches on each bridge. The front part of the head
203
www.Ebook777.com
and jaws are bright yellow In life, with a red spot on the top of the
snout; the hinder part of the head and the neck are brown. The
limbs and tail are black. This species is said to retreat to short bur
rows underground and to be entirely terrestrial. In captivity the
type specimens fed entirely on vegetable food.
Heosemys depressa is only known from the hills of Arakan.
The carapace, which reaches a length of about ten inches, is very
depressed, with a prominent median keel, serrated hind margin,
and slightly reverted front and hind margins. The head is of mode
rate size, the upper jaw bicuspid, and the skin on the hinder part of
the head is divided into large scales. The scalation of the front
limbs is similar to that of H. silvatica. The hind limbs are flattened
and are very large in adult males. The fingers are moderately
webbed. The carapace is light brown in color, uniform or with dark
mottlings; the plastron is yellowish-brown with dark blotches or
broad rays on each scute. The habits of this species are unknown.
Heosemys leytensis is a rare species only recorded from the
island of Leyte in the Philippines. The type specimen, an adult
male, is about 8 inches long and has a rather narrow, unkeeled
carapace with a slightly serrate hind margin and rather strongly
serrate front margin. The reddish-brown plastron is strongly con
stricted between the gular and humeral scutes. The head is large
and covered with smooth skin which is divided into scale-like ele
ments posteriorly. A narrow transverse yellow line crosses the
posterior part of the head and continues to the posterior border of
the tympanum. The upper jaw is slightly bicuspid and has a dis
tinct hook. Only one other specimen of this species has been de
scribed ; this differs from the type in being much smaller and in
having a slight keel on the posterior part of the carapace, in show
ing distinct growth annuli on the carapace, and in having a yellow
plastron and a yellow spot on each side of the lower jaw.
GENUS MELANOCHELYS
The genus Melanochelys includes two species, M. tricarinata
and M. trijuga. They differ from Heosemys in having tricarinate
shells and usually complete bony arches in the cheeks. Melano
chelys tricarinata is a terrestrial hill species known from Chota
Nagpur, northern Bangladesh, and the Daflas Hills and Bisnath
Plain in northern Assam. The carapace is about six inches long in
adults and of a dark plum color with three well-defined keels out
lined in dark brown. The shell is elongate and rather strongly
arched. The plastron is yellowish-brown. The head is of moderate
size, with a feeble median notch in the upper jaw. There are en
larged squarish or pointed scales on the front face of the limbs.
The head and limbs are dark plum or blackish above, with a broad
red stripe on each side of the head, starting from the nostril and
204
passing above the eye and the tympanum; a similar band passes
below the angle of the mouth along the lower margin of the lower
jaw. The limbs are sometimes decorated with yellow spots. Large
females have slightly kinetic hind plastral lobes which may assist
in oviposition as in the American Rhinoclemys.
Melanochelys trijuga is distinguished from M. tricarinata by
the entirely, or almost entirely, black or brown plastron. The cara
pace is also completely black (or dark brown in some young speci
mens) and is tricarinate in all except very old individuals. The six
subspecies extend from Ceylon north through India to Burma; in
most cases the ranges of the different races do not come in contact.
M. trijuga trijuga is found in the central part of the peninsula
of India, reaching both coasts and extending as far north as Bom
bay on the west coast. The carapace is moderately depressed, tri
carinate, and the lateral marginals are somewhat reverted. The
overall coloration of the shell is dark brown or blackish, while the
plastron has a yellow margin which diminishes in width with age.
The lateral keels in the young are lighter in color than the rest of
the shell. The skin of the head is greyish or olivaceous with yellow
or pink reticulations, best marked on the sides. The head is of
moderate size, and the upper jaw is mesially notched. The toes are
more or less completely webbed. A carapace length of about nine
inches is reached.
M.t. edeniana from Burma (Arakan; Karenni Foothills; Moul-
mein) is very similar to the typical race but differs in the larger
adult size (over eleven inches). The head is grey or brown with in
distinct yellowish reticulations which disappear at an early age.
An orange-brown marking on the lower jaw extends, with inter
ruptions, as far back as the tympanum. The ranges of trijuga and
edeniana are separated by the range of a third subspecies, indo-
peninsularis, which is found in Chota Nagpur and the Jalpaiguri
District of northern Bangladesh. Again, the most obvious differ
ence is in the adult size; the maximum length of this race is about
thirteen inches. The upper part of the head is light yellow-buff in
color, with a longitudinal black stripe that is fairly broad at the
nostrils but tapers evenly to a point at the back of the head. The
carapace is black with yellow markings on the periphery.
The largest subspecies is Melanochelys trijuga parkeri, which
reaches a carapace length of about fifteen inches. The carapace
scutes are frequently a dark cuprous-red, but this is a superficial
encrustation which may be scraped off to reveal a uniform black.
The plastron is dark brown, usually with a diffuse, dirty yellow
lateral margin. The head of the adult is either a uniform
olive-brown or is finely spotted with orange. The keels are not dis
tinct in large specimens. It is mainly vegetarian and is found both
on land and in still water. It is confined to Sri Lanka.
205
www.Ebook777.com
Melanochelys trijuga thermalis is the common ‘hard-shalled
terrapin’ of Sri Lanka and is also found in the Maidive Islands and
the Ramnad District of mainland India. Adults reach a length of
eight or nine inches. Both carapace and plastron are black, with a
slight reduction in pigmentation along the sides of the plastron and
along the three dorsal keels. The front faces of the forelimbs, the
upper faces of the hind limbs, and the top of the head are black,
other parts of the body being greyish to almost white. The head is
marked with numerous yellow spots in young specimens, but these
completely disappear as the turtle matures. The carapace is at
tractively sculptured both with concentric annuli and with radia
ting striae on each scute. In diet this turtle is omnivorous. The eggs
number from three to eight and are about 1.9 inches by 1 inch in
size. Egg laying apparently takes place throughout the year, and
one female probably lays several times during one year. This tur
tle is found both on land and in water with comparable frequency.
Newly-captured specimens, or rarely-handled captives, give off a
strong and distinctive odor when handled.
Melanochelys trijuga coronata is found in southwestern India
from Travancore and Cochin north to Calicut. It is a small race,
not exceeding a carapace length of about eight inches, and has
both carapace and plastron dark brown or black without light
markings. This race is easily identified by the head pattern, in
which the yellow temporal regions are separated by a broad dia
mond-shaped black marking which begins at the snout, widens
steadily to a point behind the eyes, and then tapers evenly to end at
the back of the head; this black marking is much broader than that
of indopeninsularis.
GENUS CYCLEMYS
A turtle which has a close superficial resemblance to Geoemy-
da spengleri is Cyclemys mouhoti, from Laos, Viet Nam, and Hai
nan Island. McDowell places the two forms in the same genus, but
a more usual position, emphasizing horizontal rather than vertical
relationships, places the species in different genera on the basis of
the development of a hinged plastron in mouhoti, though this is
only evident in mature specimens. Other authors place mouhoti in
a separate genus altogether (Pyxidea). The carapace is brown,
moderately elevated, tricarinate, and strikingly flat-topped. The
hind margin is strongly serrate, and the front margin is feebly ser
rate in hatchlings and in old specimens. A length of about seven
inches is reached. The plastron is smaller than the opening of the
shell and is yellowish-brown in color, usually with a dark spot on
each shield. The head is of moderate size, the snout truncated, and
the upper jaw strongly hooked. The skin of the hind part of the
206
Three subspecies of Melanochelys trijuga:
M. trijuga cor-
onata. Specimen
from Columbus
Zoo, Ohio.
M. trijuga ede-
niana. Photo by
Wayne Frair.
M. trijuga trijuga.
207
www.Ebook777.com
head is divided into quite large shields. The scalation of both fore
and hind limbs is heavy. The digits are partially webbed.
The other species of Cyclemys, C. dentata, is a much more
familiar form which is basically a flattened, wide-shelled version
of mouhoti. It is found in Burma, Malaya, Sumatra, Java, Nias,
the Natuna Islands, Banka, Thailand, northern India, Viet Nam,
Borneo, and the islands of Palawan, Balabac, and Leyte, in the
Philippines. No subspecies are considered valid, but the coloration
of this species is very variable. A typical specimen has an almost
uniform brown shell, serrated behind and with a slight median
keel. The plastron is of a similar color, but with narrow black radi
ating lines on each scute. The soft parts are mostly grey mottled
with yellow or pinkish orange; these mottlings form definite
stripes on the neck. The upper jaw is hooked, and the top of the
head is uniform brown. Many of the scales on the front face of the
forelimbs are wide and band-like. A hinge develops in the plastron
in mature specimens (the mature length is 9 or 10 inches), but even
in specimens six or seven inches long there is virtually no sign of
hinge development and the strong angle at which the seams
between the pectorals and abdominals meet on the midline gives
the impression that no hinge could develop at that site. However,
as maturity is approached, the bridge becomes invaded by flexible
tissue and a hinge develops right across the front part of the
abdominal scutes.
Cyclemys mouhoti from Southeast Asia is often assigned to the genus Pyx-
idea. A plastral hinge is developed only in fully mature specimens.
Plastral views of Cyclemys mouhoti (left) and Cyclemys dentata (right). The
plastral hinge can be seen in the left photo. Photo of C. mouhoti by Dr. Robert
Mertens.
Young specimens of Cyclemys dentata are sometimes con

fused with Heosemys grandis. However, grandis has more black
pigmentation on the underside, the radiating streaks being
broader and the areolae completely black, as well as a deeper
shell, a conspicuously bicuspid upper jaw, and the same mottled
pattern on the top as on the sides of the head. Cyclemys dentata is
a responsive and hardy pet turtle and will eat both vegetable and
animal food, in water or on land. In the wild state young specimens
are aquatic but adults are almost entirely terrestrial. Two to four
eggs, which measure 2.25 x 1.25, inches, are laid at a time.
R. Bourret in 1939 described a new species of emydid turtle
which he named Geoemyda tcheponensis, based upon a single
juvenile specimen, 2.75 inches long, from Se-Bang-Hien, in the
mountainous area of central Viet Nam. Bourret reported a further
specimen in 1941 from Dong-Tam-Ve. Wermuth and Mertens, in
their 1961 checklist, considered the species valid, but S.B. Mc
Dowell, in his 1964 revision of the Emydidae, suggested that Bour-
ret’s type specimen was probably a Cyclemys dentata with the
entoplastron abnormally far forward. The form is certainly super
ficially similar to Cyclemys dentata, though whether tcheponen-
209
www.Ebook777.com
sis, which is known only from juveniles, develops a plastral hinge
with maturity as does Cyclemys remains uncertain. However, sev
eral juvenile turtles imported into California in the last few years,
which were brought to my attention by Robert McMorris of the San
Francisco Zoo, appear to be indistinguishable from Bourret’s tche-
ponensis and different in a number of subtle but constant features
from dentata. I would suggest therefore that tcheponensis be con
sidered valid, and be provisionally placed in the genus Cyclemys
pending the availability of mature specimens.
The carapace of Cyclemys tcheponensis is slightly narrower
than that of juvenile Cyclemys dentata, may be slightly flared pos
teriorly, is serrate behind, has a well-defined median keel, and
tends to be rather strongly declivous at the level of the junction of
the fourth and fifth central scutes. In color the carapace is brown,
with dark brown to black areolae and with traces of both annuli
and radiating striae on each scute. The plastral scutes bear pat
terns of fine radiating black lines on a brown background. The soft
parts are similar to those of Cyclemys dentata, except that the top
of the head is somewhat mottled and there is a prominent pinkish-
buff line extending back from behind the eye along the neck. The
Specimen agreeing well with the description of Cyclemys tcheponensis, ten

tatively considered distinct from C. dentata.
210
tail is of medium length, thin and pointed. The snout is somewhat

conical, with the nostrils well anterior to the mouth, and the upper
jaw has a slight median indentation. Small axillary and inguinal
scutes are present. The digits are short but are completely web
bed.
GENUS CUORA
The Asiatic box turtles of the genus Cuora are generally con
sidered closely related to Cyclemys, though in skull features as
well as in external appearance of the juveniles Cuora appears
more similar to Mauremys, particularly Mauremys mutica. The
genus is readily identified by the well-developed plastral hinge,
present even in juveniles, which in three of the five species allows
the shell to be closed up as tightly as in the American box turtles
(Terrapene). The postorbital bar is present in some species of this
genus, absent in others.
The two apparently more primitive species are C. trifasciata
and C. yunnanensis, in which the carapace is rather low and the
hind lobe of the plastron too narrow to close the shell opening tight
ly. In both of these species there is a well-defined notch between
the anal scutes.
Cuora trifasciata occurs in southern China (Kwang Si and
Kwang Tung Provinces), on Hainan Island, and in northern Indo-
Plastral view of Cuora am-

boinensis. Compare the hinge
with that of Terrapene. Photo
by G. Marcuse.
211
www.Ebook777.com
China. This species has an elongate, moderately depressed cara
pace, the widest point of which is well behind the middle. The cara
pace is brown, with three longitudinal keels outlined in black; the
median black stripe extends from the middle of the first central
right to the rear of the shell, but the lateral stripes are only present
on the first three costals on each side. The plastron is almost
entirely black except for an irregular yellow border that may dis
appear on the gulars and anals and sometimes a slight lightening
of the intensity of the black pigment along the midline. The under
side of the marginals is pinkish-yellow with small black blotches
present on the posterior border of some of the anterior and lateral
marginals. The upper jaw is feebly hooked; the top of the head is
immaculate yellow or olive-yellow. A black streak passes along
each side of the head, including the eye, and encloses two oval yel
low spots, the larger immediately behind the eye. A second thinner
black streak extends back from the border of the mouth. The soft
skin is light pink in color, the limbs pink vermiculated with grey. A
shell length of about eight inches is reached. The species has been
found in clear mountain streams in southern Kwang Tung, at be
tween 150 and 1300 feet altitude. Oviposition probably takes place
in May, and two eggs are laid at a time, measuring about two
inches by one inch. The species does well in captivity but is exceed
ingly shy even after long periods in confinement, so that the attrac
tive head markings can rarely be seen.
Cuora yunnanensis is closely related to trifasciata but differs
in having a shorter snout, unhooked upper jaw, and an olive-tinged
plastron with a yellowish border and blackish seams. The soft
parts are marbled with orange instead of pink. This species is
found at much higher altitudes than trifasciata; the two type local
ities (Yunnan Fu and Tong Chuan Fu, Yunnan Province, China)
are about 7000 feet above sea level.
The Amboina or Malayan box turtle, Cuora amboinensis, is a
well-known and abundant species from Tenasserim, Thailand,
Cambodia, Viet Nam, Malaysia, Indonesia as far east as Sulawesi
and Amboina, and Luzon, Polillo, Dinagat, and Mindanao in the
Philippines. The carapace reaches a length of about eight inches,
is moderately domed, has a median keel and sometimes two
lateral keels (especially in the young), and is dark olive, brown, or
black in color. The plastron is yellow or light horn color, usually
with a dark blotch on the outer and posterior part of each scute,
and there is often a dark blotch on the postero-lateral comer of
some or all of the under-marginals. The skin of the head is smooth,
the upper jaw feebly hooked, and the toes fully webbed. The head
is dark brown or green above, yellow below, and with a bright yel
low band bordering the top of the head on each side, the two bands
meeting above the nostrils. A second paler band passes through
the eye and is separated from the margin of the upper jaw by a
212
dark band; a third borders the mandible. The Malayan box turtle
Is more aquatic than its American counterparts, inhabiting ponds,
marshes, and flooded rice paddies, though on occasion it is found a
long way from water. Malcolm Smith reports that it is entirely
vegetarian and only eats small quantities at a time; however, in
captivity it becomes omnivorous and quite voracious. The eggs
measure about 1.75” x 1.25”.
Cuora galbinifrons is only known from Bach-Ma, Tam-Dao,
and Linh-Can, Viet Nam. The carapace is less domed than that of
amboinensis, dark brown in color, and with a strong yellow verte
bral stripe and less distinct costal stripes. The areolae of the mar
ginal scutes are also yellow. The plastron is almost uniform black,
but the entire submarginal area as well as a limited area on the
edges of the pectorals and abdominals is orange-yellow. The upper
jaw is strongly hooked, and the head is pale olive-green on the
sides, darker on top. A narrow, black-bordered white line extends
from the nostril to the eye, and this line is continued behind the eye
as a much broader and stronger white line, again with a black bor
der. The habits of this species are unknown.
Cuora flavomarginata is known from southern China (Tung-
ting-See), Taiwan, and Ishigaki Shima and Iriomote in the Ryu-
Kyu Islands. This species reaches a length of about seven inches
and has a dark brown or black carapace, the areola of each scute
being a lighter brown. A median keel is present in all except old in
dividuals, and each scute is marked with well-sculptured growth
annuli. The plastron is similar to that of C. galbinifrons— black, or
dark brown, with yellow under the marginals and on the outer
parts of the pectorals and abdominals. The top of the head is olive-
brown, the sides and underneath yellow, with a light, dark-edged
stripe through the eye that again is very similar to that of galbini
frons. Malcolm Smith reports that this species is found in ponds
and flooded rice fields, but its habits have not been studied.
GENUS HIEREMYS
The genus Hieremys, with its single species, annandalei, is
closely related to Geoemyda, from which it differs in having broad
jaw surfaces which lack a median ridge, and with a corresponding
posterior displacement of the bony choanae (which are below the
middle of the orbits in Hieremys, below the anterior part of the
orbits in Geoemyda). Hieremys annandalei is found in central
Thailand, Cambodia, Viet Nam, and the northern part of the
Malay Peninsula. It is found not only in slow-moving rivers right
down to the seacoast, but also in swamps and marshy fields. This
is a large species, a shell length of eighteen inches being reached.
The carapace is rather low, with a median keel, and uniform dark
brown or black in color. The plastron is unmarked yellow in young
213
www.Ebook777.com
Head and plastron of Hieremys annandalei. Specimen from the Lantz collec-
tion.
214
specimens, but with growth a black patch develops on each plas-

tral scute, which may eventually extend to cover the entire scute.
The soft parts are greenish or greyish; young specimens have a
prominent yellow stripe passing from each nostril, above and
through the eye, and extending backward above the tympanum to
the neck, with other yellow stripes along the upper and lower jaws
and along the neck. These streaks usually disappear in old individ
uals, in which the head becomes greyish with yellow or green ver-
miculations. This is a herbivorous species. It is frequently cap
tured and kept as a pet in Malaya and is said to be readily tamed.
This is one of the turtles often deposited in the tanks and canals
around the Tortoise Temple in Bangkok, for reasons connected
with the Buddhist belief that ‘saving’ a turtle in this way gains one
merit in the next world.
GENUS NOTOCHELYS
Notochelys platynota, found in South Viet Nam, the Malay
Peninsula, Borneo, and Sumatra, is a moderately large species
that may be related to Cyclemys. The carapace, which may reach
a length of thirteen inches, is rather depressed, strongly serrated
behind, and flat-topped; the back of the shell falls off at a sharp
angle toward the rear just behind the fourth central; at the actual
site of this angle, a small square extra central scute is located,
intercalated between the homologs of the fourth and fifth centrals
in other turtles. This extra central scute is present in other turtles
only as an abnormality (for example in three out of 24 Hieremys
annandalei examined by Malcolm Smith), and this is the only spe-
Notochelys
platynota. Notice
the extra central
scute. Specimen
from the Colum
bus Zoo, Ohio.
215
www.Ebook777.com
cies in which it is constant. The carapace of hatchling Notochelys
is a most attractive apple-green color, with a pair of black spots on
each central scute, and sometimes a single spot on each costal as
well. With maturity, these spots disappear and the shell assumes a
red-brown color. The underside is orange-red at all ages, but in
adults extensive areas of dark pigment are deposited on the plas
tron and bridge. A feeble mid-plastral hinge is present in large in
dividuals. In young specimens a broad irregular yellow V-shaped
mark is present on top of the head, the apex being above the nos
trils and the rami passing above the eyes. Another yellow line ex
tends backward from the comer of the mouth on each side. These
markings become faint or disappear with age. The upper jaw is
strongly hooked. The toes are fully webbed, and the front face of
the forelimbs and back of the hind limbs bear transversely en
larged scales. This species is said to inhabit ponds and swamped
jungles and to eat vegetables and fruit. In captivity, even very
young specimens live happily on an entirely vegetarian diet. On
the other hand, Stanley Flower noticed two specimens attacking a
large, sluggish water snake (Achrochordus javanicus). This is
said to be one of the most active and conspicuous of the Malayan
turtles, but its breeding habits have not been described.
GENUS OCADIA
Ocadia sinensis is a fairly small species which is abundant on
Hainan and Taiwan but apparently rare on the mainland of China,
though Arthur Stanley reports it from Fukien, Hangchow, Soo-
chow, and Shanghai in southern China. It is also known from North
Viet Nam. Loveridge and Williams consider this species and Pseu-
demys to be among the most primitive of living emydids. Structur
al features which identify this turtle are the entoplastron inter
sected by the humero-pectoral suture, the long tail, the fully web
bed digits, the presence of only three phalanges in the fifth finger
and fifth toe, and the bony choanae on a level with the middle of the
eyes. The carapace, which may reach a length of nearly ten
inches, is moderately domed and elongate, tricarinate in young
specimens, and brown to black in color; young specimens have
orange markings on each of the costal and central scutes. The
underside is yellow, each shield with a large brown blotch, the pec
torals and abdominals with two. In Taiwan specimens the brown
blotches may be very pale or absent. The head, neck, limbs,
thighs, and tail are all marked with numerous narrow, pale green
longitudinal stripes; these stripes are most apparent on the sides
of the head. The upper jaw is white, and the iris of the eye is pale
green, with a black bar running through the pupil similar to that of
many American emydids. This species is reported to inhabit quiet
water (ponds, canals, and sluggish streams) in open country at
low elevations. In captivity is prefers vegetable to animal food.
216
One captive specimen reported by Malcolm Smith laid three eggs

approximately 1.6” x 1” in size. Ocadia sinensis is eaten by the
natives of Hainan.
GENUS MORENIA
The genus Morenia includes two species, M. ocellata, known
from southern Burma and from Toungoo to Mergui in Tenasserim,
and Ikf. petersi, known from the Jessore District, Dacca, and
Fatehgarh in Bangladesh. The genus is characterized by the fail
ure of the humero-pectoral suture to traverse the entoplastron, the
posteriorly-placed choanae (behind the level of the eyes), and the
divided skin on the hinder part of the head. In both species the car
apace is moderately domed (more so in females than in males),
the upper jaw is mesially notched, the jaw surfaces are broad and
ridged, and the posterior margin of the carapace not serrated.
Morenia ocellata reaches a shell length of nine inches. The cara
pace is light or dark olive-brown in color, with a large, pale, yellow
ocellus on each scute, with a dark brown center and less distinct
The dark-centered
ocelli of Morenia
ocellata are
distinctive.
Specimen from
British Museum
(Natural History).
dark border. The underside is immaculate yellow. The ground

color of the head is olive or brownish, with a pale yellow streak on
each side passing from the snout above the eye on to the neck and
another from behind the eye to above the tympanum. This species
is reportedly strictly aquatic, but Theobold reports that large
numbers are often left dry by the drying up of inundated plains
during hot weather; in such situations many are captured for food
by the Burmese. This species is very rarely imported into the
United States, and its habits in the wild or in captivity have not
been recorded.
217
www.Ebook777.com
Morenia petersi is distinguished from M. ocellata by the much
longer snout and the wider nuchal scute. The carapace is olive or
black in color, the vertebral and costal scutes margined with pale
green or yellow. Females reach a length of about eight inches;
males are smaller. The first four vertebrals each bear a narrow,
pale green longitudinal stripe and a U-shaped mark with the ends
directed forwards. Each costal and the last vertebral has a large
pale green circle placed low down, with irregular looped lines of
the same color above. Each marginal scute is decorated with a
vertical pale green streak. The underside is yellow or orange, with
a dark blotch on each axillary scute and the undersides of the
fourth, fifth, and sixth marginals. The top and sides of the head are
very dark olive with three yellow streaks, one bordering the top of
the head, one behind the eye, and one below the eye starting from
the tip of the snout. The limbs are bordered with yellow.
GENUS GEOCLEMYS
The black pond turtle, Geoclemys hamiltoni, is a fairly large
species known from the Ganges and Indus River systems in India
and Pakistan. The carapace is somewhat elongate, moderately
elevated, and with three well-defined keels. The upper part of the
shell is jet black, but there are wedge-shaped dull yellow markings
The Indian black pond turtle, Geoclemys hamiltoni.
218
on the lower part of the costals and numerous dull yellow blotches
on the marginals. The carapace reaches a length of about fourteen
inches. The plastron is pale yellow with linear dark markings con
centrated on the pectoral and abdominal scutes. A very deep notch
is present between the anal scutes. The head is very large, black in
color, with large symmetrical pale yellow spots concentrated
around the snout, lower jaw, and behind the eye. The jaw surfaces
are very broad. The neck is grey with numerous white spots; the
soft skin is pink or white, but the forelimbs are black with white
spots. The front faces of the forelimbs are covered with wide,
band-like scales. This species is recorded from oxbow lakes and
sloughs in quiet, shallow, rather clear water with considerable
aquatic vegetation. The diet in the wild appears to consist princi
pally of snails; in captivity meat and fish are also accepted, but
not vegetable matter. The reproductive habits of this species have
not been recorded. The young are much more brightly colored
than the adults.
GENUS CHINEMYS
The well-known Reeves’ turtle, Chinemys reevesi, used to be
placed in the genus Geoclemys but was transferred to a new genus
because of its longer tail, more posteriorly placed entoplastron,
and more anteriorly located choanae (level with the middle of the
eyes). Chinemys reevesi is found in southern China through the
Yangtze Valley from the junction of the Yangtze with the Min east
ward to the sea and along the Chinese seaboard from northern
Kiangsu or southern Shantung, at least as far as Canton, and is
known from the Wei Valley in Shensi. The species is also known
from Korea and southern Japan, but not from Hainan. This species
is of considerable economic importance and has been introduced
to many places outside its natural range. It is generally considered
to be a very small species, and it is true that Chinese specimens
only four or five inches long show well-defined sexual characteris
tics and no evidence of recent growth. On the other hand, Sam Tel
ford informs me that Japanese specimens may get to be well over
a foot in length and possibly fifteen inches. The carapace is paral
lel-sided and moderately domed, with three strong keels. The
carapace scutes are brown, edged with yellow or white. The plas
tron is strongly notched behind and is basically yellow in color, but
large black blotches on each plastral scute may reach almost to
the seams. The head is of moderate size, the upper jaw sometimes
notched mesially, the digits are fully webbed, and the tail is long.
The head is greyish-olive in color, with yellow dark-edged longitu
dinal streaks and angular markings. The limbs are grey, some
times with yellow spots. Reeves’ turtle is omnivorous in diet and is
tough and hardy in captivity. It is very abundant in some parts of
219
www.Ebook777.com
its range, especially the lower Yangtze Valley. During courtship,
the males are reported to parade around the female in a tireless
effort to rub their snouts on hers. Four to six eggs are laid at a
time.
Quite frequently, specimens of Chinemys reevesi are found in
which the entire shell and soft parts are very dark brown or black.
220
This phenotype has received a varietal name (unicolor), and it

seems to occur quite frequently in Japanese specimens; but since
it may occur anywhere in the range of the species, it must be con
sidered simply as a melanistic variant rather than a subspecies.
The species Chinemys grangeri (originally described as Geo-
clemys grangeri by Karl Schmidt in 1927) is customarily synony-
mized with reevesi, the type specimen being the only preserved
example known and showing a general resemblance to reevesi.
However, Robert McMorris of the San Francisco Zoo recently in
formed me that a turtle imported in small numbers to the United
States in the last year or so from northern Taiwan may in fact be
referable to grangeri, which he considers a valid species. G. gran
geri was originally based upon a 4.3” male specimen from Yen-
chingkao, Wanhsien, Szechwan, China, at 1500 feet altitude; thus,
the enormous distance between Taiwan and the type locality of
grangeri would argue against the two populations being identical.
The type of grangeri was distinguished from reevesi by having the
An adult Chinemys reevesi. The head pattern is poorly developed. Photo by

G. Marcuse.
221
www.Ebook777.com
Specimen of Chinemys reevesi that agrees with the tentatively valid
Chinemys grangeri.
axillary scute larger than the inguinal, the small occipital shields
much smaller, and the spots on the plastron smaller and more con
trasting. The single specimen from Taiwan which I have examined
(and which McMorris assures me is absolutely typical of the origi
nal series of six that passed through his hands) fits the description
of grangeri reasonably well, though the postocular stripe gives off
a downward process at a short distance behind the eye instead of
immediately behind it as in the type of grangeri. The general ap
pearance of the animal with head retracted is of a turtle almost
identical to Ocadia sinensis, but the different neck striping is evi
dent as soon as the head is extended.
Live animal exporters in Taiwan recognize this turtle as being
distinct from Ocadia sinensis. They report that it is confined to the
northern part of the island, where extensive pesticide spraying of
the cultivated areas has reduced the populations significantly.
The other two species of Chinemys are C. kwangtungensis and
C. megalocephala. C. kwangtungensis is only known from Lofao-
shan, Tinghushan, and Hsiaokeng, in Kwang Tung, China. It dif-
222
fers from reevesi in having wider jaw surfaces, a concave median

anterior upper jaw profile, granular skin on the back of the head
(polygonal scales in reevesi), a single carapace keel, and relative
ly inconspicuous vermiculation on the sides of the head. It lives in
mountain streams and is reported to lay only two eggs at a time.
Chinemys megalocephala is known from Nanking Province,
China. It has a much broader head than reevesi and broader crush
ing jaw surfaces. The carapace has three dark longitudinal stripes
but no keels.
GENUS MALAYEMYS
Malayemys subtrijuga, the Malayan snail-eating turtle, is
found in south Viet Nam, Thailand, and from the northern part of
the Malay Peninsula; it is also recorded from Java. The carapace
is brown, with some variation in shade, and there are three strong
keels, though the lateral ones do not usually extend beyond the
third costal. The hind margin of the carapace is not serrated. The
A juvenile Malayan snail-eating turtle, Malayemys subtrijuga, from Southeast

Asia. Photo by R.J. Church.
223
www.Ebook777.com
plastron is mainly dark brown or black, but with a narrow yellow
rim to each scute. The head is large, black in color, with a broad,
projecting snout and strong white lines running backward, above
the eyes, on each side, starting at the snout. Other white lines run
from the snout below the eye and along the jaw,.and a shorter line
extends backward from the middle of the eye on each side. The
upper jaw is emarginate mesially, and the digits are webbed. The
soft parts are brown. This species inhabits canals, slow-moving
rivers, and marshy places. It subsists mainly on snails, but occa
sionally takes other animal food. Most specimens imported into
the United States are only a few inches long, but large adults are
well over a foot in length. The eggs are about 1.75 inches x 1 inch in
size.
GENUS SIEBENROCKIELLA
Siebenrockiella crassicollis is another species which is nor
mally considered to be very small but which may exceed fifteen
inches in carapace length. The shell is moderately depressed and
in half-grown and adult specimens only shows a central keel,
though lateral keels may be present in young specimens. The over
all color is dark brown or black, but the plastral seams tend to be
of a lighter color and there are white spots on the head—one above
and in front of the eye on each side and several others behind the
angle of the mouth. The head is moderately large, and the digits
are fully webbed. There is a strip of granular skin from the eye to
the tympanum. This species is found in Tenasserim, Thailand,
South Viet Nam, the Malay Peninsula, and Sumatra, Borneo, and
Java. It lives in ponds, marshes, and sluggish streams; it is entire
ly carnivorous, feeding on molluscs, worms, frogs, and offal.
Detail of head of
Siebenrockiella crassicollis.
224
A juvenile red-eared slider, Pseudemys scripts elegans, from DeKalb County,

Georgia.
www.Ebook777.com
BACKGROUND READING
EMYDIDAE - GENERAL:
McDowell, S. B. 1964. Partition of the genus Clemmys and
related problems in the taxonomy of the aquatic Testudini-
dae. Proc. Zool. Soc. London, 143: 239-279.
UNITED STATES SPECIES:
Carr, A.F. 1952. Handbook of turtles. Comstock Publ. Asso
ciates, Cornell Univ. Press. 542 pp.
Conant, R. 1958. A field guide to reptiles and amphibians.
Houghton Mifflin Co., Boston. 366 pp.
Ernst, C.H. and R.W. Barbour. 1972. Turtles of the United
States. Univ. of Kentucky Press. 347 pp.
Pope, C.H. 1939. Turtles of the United States and Canada.
Alfred A. Knopf, New York. 343 pp.
ASIATIC SPECIES:
Pope, C.H. 1935. The reptiles of China. Natural History of Cen
tral Asia, Vol. X. Amer. Mus. Nat. Hist., N.Y. 604 pp.
Smith, M.A. 1931. The fauna of British India, including Ceylon
and Burma. Reptilia and Amphibia. Vol. 1. Loricata, Testu-
dines. Taylor and Francis, London. 185 pp.
PSEUDEMYS:
Alvarez del Torro, M. 1960. Los reptiles de Chiapas. Inst. Zool.
del Estado, Textla Gutierrez, Chiapas, Mexico. 204 pp.
Barbour, T. and A.F. Carr. 1940. Antillean terrapins. Mem.
Mus. Comp. Zool. 54(4): 381-413.
Cagle, F.R. 1937. Egglaying habits of the slider turtle
(.Pseudemys scripta troosti), the painted turtle (Chrysemys
picta), and the musk turtle (Stemotherus odoratus). J.
Tennessee Acad. Sci., 12: 87-95.
__________ 1944a. Activity and winter changes of hatchling
Pseudemys. Copeia 1944: 105-109.
__________. 1944b. Home range, homing behavior and migra
tion in turtles. Misc. Publ. Mus. Zool. Univ. Michigan 61:
1-34.
__________. 1944c. Sexual maturity of the female of the turtle
Pseudemys scripta elegans. Copeia 1944: 149-152.
__________ 1946. The growth of the slider turtle Pseudemys
scripta elegans. Amer. Midi. Nat. 36: 685-729.
________ 1947. Color abnormalities in Pseudemys scripta
troostii (Holbrook). Nat. Hist. Misc. 6: 1-3.
__________ 1950. The life history of the slider turtle, Pseudemys
scripta troostii (Holbrook). Ecol. Monogr. 20 : 31-54.
Carr, A.F. 1935. The identity and status of two turtles of the
genus Pseudemys. Copeia 1935: 147-148.
______ . 1937a. A new turtle from Florida, with notes on
226
Pseudemys floridana mobiliensis (Holbrook). Occ. Pap.

Mus. Zool. Univ. Michigan 348: 1-7.
------------- 1937b. The status of Pseudemys scri-pta (Schoepff)
and Pseudemys troostii (Holbrook). Herpetologica 1: 75-77.
------------- 1938a. Pseudemys nelsoni, a new turtle from Flori
da. Occ. Pap. Boston Soc. Nat. Hist. 8: 305-310.
------------- 1938b. A new subspecies of Pseudemys floridana,
with notes on the floridana complex. Copeia 1938: 105-109.
------------- 1938c. Notes on the Pseudemys scripta complex.
Herpetologica 1: 131-135.
------------- 1942. The status of Pseudemys floridana texana with
notes on parallelism in Pseudemys. Proc. New England
Zool. Club 21: 69-76.
__________ 1952. op. clt.
------------ and J.W. Crenshaw. 1957. A taxonomic reappraisal
of the turtle Pseudemys alabamensis Baur. Bull. Florida
State Mus. 2: 25-42.
Casas-Andreu, G. 1967. Contribucion al conocimiento de las tor-
tugas dulceacuicolas de Mexico. Tesis Professional, Biol.
Dept., Univ. Nac. Aut. de Mexico, 96 pp.
Freiberg, M.A. 1969. Una nueva subespecie de Pseudemys dor-
bignyi Dumeril et Bibron) (Reptilia, Chelonia, Emydidae).
Physis, Buenos Aires, XXVIII, 77: 299-314.
-------------1971. El mundo de las tortugas. Editorial Albatros,
Buenos Aires. 134 pp.
Gibbons, J.W. 1969. Structure and dynamics of the yellow-bel
lied turtle, Pseudemys scripta (Reptilia: Chelonia:
Emydidae) in an artificially heated reservoir. A.S.B. Bull.,
16: 52.
------------- 1970. Reproductive dynamics of a turtle (Pseu
demys scripta) population in a reservoir receiving heated
effluent from a nuclear reactor. Canadian J. Zool. 48: 881-
885.
Grant, C. 1940. The herpetology of the Cayman Islands. Inst.
Jamaica, 65 pp.
Legler, J.M. 1960a. A new subspecies of slider turtle
(Pseudemys scripta) from Coahuila, Mexico. Univ. Kansas
Publ., Mus. Nat. Hist. 13(3): 73-84.
------------- 1960b. Remarks on the natural history of the Big
Bend slider, Pseudemys scripta gaigeae Hartweg. Herpeto
logica 16(2): 139-140.
------------ and R.G. Webb. 1970. A new slider turtle (Pseu
demys scripta) from Sonora, Mexico. Herpetologica 26(2):
157-168.
Medem, F. 1962. La distribucion geographica y ecologia de los
Crocodylia y Testudinata en el Departamento del Choco.
227
www.Ebook777.com
A young Cumberland turtle, Pseudemys scripta troosti, a subspecies from
Tennessee and Kentucky.
228
Head and plastral views of the Baja California cooler, Pseudemys scripts
nebulosa.
www.Ebook777.com
Rev. Acad. Colombiana Cien. Exactas, Fis. y Natur. 11(44):
279-303.
Moll, E.O. and J.M. Legler. 1971. The life history of a neotropi
cal slider turtle, Pseudemys scripta (Schoepff), in Panama.
Bull. Los Angeles Co. Mus. Nat. Hist. Sci., No. 11: 1-102.
Strecker, J.K. 1927. Observations on the food habits of Texas
amphibians and reptiles. Copeia, No. 162: 6-9.
Williams, E.E. 1956. Pseudemys scripta callirostris from Vene
zuela with a general survey of the scripta series. Bull. Mus.
Comp. Zool. Harvard 115(5): 145-160.
CHRYSEMYS:
Bleakney, J.S. 1958. Postglacial dispersal of the turtle Chry-
semys picta. Herpetologica 14: 101-104.
Cagle, F.R. 1954. Observations on the life cycles of painted tur
tles (genus Chrysemys). Amer. Midi. Nat. 52: 225-235.
Emlen, S.T. 1969. Homing ability and orientation in the painted
turtle Chrysemys picta marginata. Behaviour 33: 58-76.
Ernst, C.H. 1967. Intergradation between the painted turtles
Chrysemys picta picta and Chrysemys picta dorsalis.
Copeia 1967: 131-136.
__________ 1969. Natural history and ecology of the painted tur
tle, Chrysemys picta (Schneider). Ph.D. dissertation, Uni
versity of Kentucky. 202 pp.
__________ 1970. The status of the painted turtle Chrysemys
picta in Tennessee and Kentucky. J. Herpetol. 4: 39-45.
__________. 1971a. Sexual cycles and maturity of the turtle Chry
semys picta. Biol. Bull. 140: 191-200.
__________ 1971b. Population dynamics and activity cycles of
Chrysemys picta in southeastern Pennsylvania. J. Herpetol.
5: 216-220.
Gibbons, J.W. 1967. Variation in growth rates in three popula
tions of the painted turtle Chrysemys picta. Herpetologica
23: 296-303.
__________. 1968a. Carapacial algae in a population of the
painted turtle Chrysemys picta. Amer. Midi. Nat. 79: 517-
519.
__________ 1968b. Population structure and survivorship in the
painted turtle, Chrysemys picta. Copeia 1968: 260-268.
__________ 1968c. Reproductive potential, activity, and cycles
in the painted turtle, Chrysemys picta. Ecology 49: 399-409.
Kosh, R. J. and V.H. Hutchison. 1968. Daily rhythmicity of tem
perature tolerance in eastern painted turtles, Chrysemys
picta. Copeia 1968: 244-246.
Legler, J.M. 1954. Nesting habits of the western painted turtle,
Chrysemys picta hellii (Gray). Herpetologica 10: 137-144.
Sexton, O. J. 1959. Spatial and temporal movements of a popula
230
tion of the painted turtle, Chrysemys picta marginata

(Agassiz). Ecol. Monogr. 29: 113-140.
GRAPTEMYS:
Cagle, F.R. 1952. The status of the turtles Graptemys pulchra
Baur and Graptemys barbouri Carr and Marchand, with
notes on their natural history. Copeia, 4: 224-234.
__________ 1953. The status of Graptemys oculifera. Zoologica,
38: 137-144.
__________ 1954. Two new species of the genus Graptemys.
Tulane Stud. Zool., 1(11): 167-186.
Carr, A.F. and L. J. Marchand. 1942. A new turtle from the Chip-
ola River, Florida. Proc. New England Zool. Club, 20:
95-100.
Chaney, A. and C.L. Smith. 1950. Methods for collecting map-
turtles. Copeia 1950: 323-324.
Floyd, P. 1973. Singing river sawbacks. Int. Turt. Tort. Soc. J.
7(2): 8-10.
Folkerts, G.W. and R.H. Mount. 1969. A new subspecies of the
turtle Graptemys nigrinoda Cagle. Copeia 4: 677-682.
Timkin, R.L. 1968. Graptemys pseudogeographica in the Upper
Missouri River of the northcentral United States. J. Herpe
tol. 1: 76-82.
Wahlquist, H. 1970. Sawbacks of the Gulf Coast. Int. Turt. Tort.
Soc. J. 4(4): 10-13,28.
Webb, R.G. 1961. Observations on the life histories of turtles
(genus Pseudemys and Graptemys) in Lake Texoma, Okla
homa. Amer. Midi. Nat. 65: .193-214.
MALACLEMYS:
Belinky, G.K. 1976. The ladies of striped pants. Mass. Herp.
Soc. Review, Vol. 7 (Winter 1976): 6-13.
Bentley, P.J., W.L. Bretz, and K. Schmidt-Nielsen. 1967. Osmo
regulation in the diamondback terrapin, Malaclemys terra
pin centrata. J. Exp. Biol. 46: 161-167.
Cagle, F.R. 1952. A Louisiana terrapin population (Mala
clemys). Copeia, 1952: 74-76.
Carr, A.F. 1946. Status of the mangrove terrapin. Copeia 1946:
170-172.
Coker, R.E. 1906. The cultivation of the diamond-back terrapin.
North Carolina Geol. Survey Bull. 14: 1-69.
__________ 1920. The diamond-back terrapin: past, present and
future. Sci. Monthly 11: 171-186.
Cowan, F.B.M. 1969. Gross and microscopic anatomy of the
orbital glands of Malaclemys and other emydine turtles.
Canadian J. Zool. 47: 723-729.
Dunson, W.A. 1970. Some aspects of electrolyte and water bal-
231
www.Ebook777.com
Two views of a Mexican slider, Pseudemys scripta cataspila, from southern
Tamaulipas, Mexico.
A Mexican slider turtle, Pseudemys scripta venusta, from Veracruz.
Hatchling Costa Rican slider, Pseudemys scripta venusta.
233
www.Ebook777.com
ance in three estuarine reptiles, the diamondback terrapin,
American and “salt water” crocodiles. Comp. Biochem.
Physiol. 32: 161-174.
Hay, W.P. 1904. A revision of Malaclemmys, a genus of turtles.
Bull. U.S. Bur. Fish. 24: 1-20.
Hildebrand, S.F. 1929. Review of experiments on artificial cul
ture of diamondback terrapin. Bull. U.S. Bur. Fish. 45: 25-
70.
__________ 1932. Growth of diamond-back terrapins, size at
tained, sex ratio, and longevity. Zoologica (N.Y.) 9: 551-563.
__________and C. Hatsel. 1926. Diamondback terrapin culture at
Beaufort, N.C. U.S. Bur. Fish., Econ. Cir. 60: 1-20.
Schwartz, A. 1955. The diamondback terrapins (Malaclemys
terrapin) of peninsular Florida. Proc. Biol. Soc. Washington
68: 157-164.
DEIROCHELYS:
Campbell, H. W. 1969. The unsung chicken turtle. Int. Turt. Tort.
Soc. J. 3(5): 22-24,36.
Gibbons, J.W. 1969. Ecology and population dynamics of the
chicken turtle, Deirochelys reticularia. Copeia 1969: 669-676
__________ 1970. Terrestrial activity and the population dy
namics of aquatic turtles. Amer. Midi. Nat. 83 : 404-414.
Krefft, G. 1951. Deirochelys reticularia (Latreille), eine wenig
bekannte Schmuckschildkrote der USA. Aquar. Terrar. Z.
4: 157-160.
Schwartz, A. 1956. Geographic variation in the chicken turtle.
Fieldiana: Zool. 34: 461-503.
EMYDOIDEA :
Bleakney, J.S. 1963. Notes on the distribution and life histories
of turtles in Nova Scotia. Canadian Field-Nat. 77: 67-76.
Cahn, A.R. 1937. The turtles of Illinois. Illinois Biol. Monogr. 35:
1-218.
Evermann, B.W. and H.W. Clark. 1916. The turtles and batra-
cians of the Lake Maxinkuckee region. Proc. Indiana Acad.
Sci. 1916: 472-518.
Gibbons, J.W. 1968. Observations on the ecology and population
dynamics of the Blanding’s turtle, Emydoidea blandingi.
Canadian J. Zool. 46: 288-290.
Lagler, K.F. 1943. Food habits and economic relations of the
turtles of Michigan with special reference to fish
management. Amer. Midi. Nat. 29: 257-312.
Smith, P.W. 1961. The amphibians and reptiles of Illinois. Illi
nois Nat. Hist. Surv. Bull. 28: 1-298.
CLEMMYS:
Adler, K.K. 1961. Egg-laying in the spotted turtle, Clemmys
guttata (Schneider). Ohio J. Sci. 61: 180-182.
234
Babcock, H.L. 1919. The turtles of New England. Mem. Boston

Soc. Nat. Hist. 8: 232-431.
Barton, A.J. and J.W. Price. 1955. Our knowledge of the bog-
turtle, Clemmys muhlenbergi, surveyed and augmented.
Copeia 1955: 159-165.
Campbell, H.W. 1960. The bog turtle in Maryland. Maryland
Nat. 30: 15-16.
Conant, R. 1951. The reptiles of Ohio. Univ. Notre Dame Press,
Notre Dame, Indiana. 284 pp.
Eglis, A. 1967. Clemmys muhlenbergi, rarest of North Ameri
can turtles. Anim. Kingdom 70: 58-61.
Ernst, C.H. 1967. A mating aggregation of the spotted turtle
Clemmys guttata. Copeia 1967: 473-474.
__________. 1968. Homing ability in the spotted turtle, Clemmys
guttata (Schneider). Herpetologica 24: 77-78.
__________. 1970a. Home range of the spotted turtle, Clemmys
guttata (Schneider). Copeia 1970: 391-393.
__________. 1970b. Reproduction in Clemmys guttata. Herpeto
logica 26: 228-232.
_________ . 1971. Clemmys insculpta. Cat. Amer. Amphib. Rept.
125: 1-2.
Evenden, F.G. 1948. Distribution of the turtles of western Ore
gon. Herpetologica 4: 201-204.
Hutchison, V.H. 1963. Record of the bog turtle, Clemmys muh
lenbergi, in southwestern Virginia. Copeia 1963: 156-157.
Nemuras, K. 1966. Genus Clemmys. Int. Turt. Tort. Soc. J. 1(1):

26-27, 39, 44.
__________. 1967a. Genus Clemmys. Int. Turt. Tort. Soc. J. 1(2):
38-40.
__________ 1967b. Notes on the natural history of Clemmys
muhlenbergi. Bull. Maryland Herpetol. Soc. 3: 80-96.
_________ . 1968. Again, the spotted turtle. Int. Turt. Tort. Soc.
J. 2(3): 32-35.
_________ . 1969. Survival of the Muhlenberg. Int. Turt. Tort.
Soc. J. 3(5): 18-21.
Netting, M.G. 1936. Hibernation and migration of the spotted
turtle, Clemmys guttata. Copeia 1936: 112.
Pallas, D.C. 1960. Observations on the nesting of the wood tur
tle, Clemmys insculpta. Copeia 1960: 155-156.
Powell, C. 1967. Female sexual cycles of Chrysemys picta and
Clemrpys insculpta in Nova Scotia. Canadian Field-Nat.
81: 134-140.
Richmond, N.D. and C.J. Goin. 1938. Notes on a collection of
amphibians and reptiles from New Kent County, Virginia.
Ann. Carnegie Mus. 27: 301-310.
235
www.Ebook777.com
Hatchling South American slider turtle, Pseudemys scripts dorbignyi.
OPPOSITE: Above: Haitian slider turtle, Pseudemys decorata. Below: Adult

female Jamaican terrapin, Pseudemys terrapen.
237
www.Ebook777.com
Storer, T.I. 1930. Notes on the range and life-history of the
Pacific fresh-water turtle, Clemmys marmorata. Univ. Cali
fornia Publ. Zool. 32: 429-441.
Yerkes, R.M. 1905. The color pattern of Nanemys guttata
Schneider. Science 21: 386.
TERRAPENE:
Allard, H.A. 1935. The natural history of the box turtle. Sci.
Monthly 41: 325-338.
------------- 1948. The eastern box-turtle and its behavior. J.
Tennessee Acad. Sci. 23: 307-321.
Brown, W.S. 1974. Ecology of the aquatic box turtle, Terrapene
coahuila (Chelonia, Emydidae) in Northern Mexico. Bull.
Florida State Mus., 19(1): 1-67.
Cahn, A.R. 1937. The turtles of Illinois. Illinois Biol. Monogr. 35:
1-218.
Carpenter, C.C. 1957. Hibernation, hibernacula and associated
behavior of the three-toed box turtle (Terrapene Carolina tri
unguis). Copeia 1957: 278-282.
Cook, S., D. Abb and W. Frair. 1972. A new record size box tur
tle. Int. Turt. Tort. Soc. J. 6(3): 1, 9-17.
Ditmars, R.L. 1934. A review of the box turtles. Zoologica
(N.Y.) 17: 1-44.
Graham, T.E. and V.H. Hutchison. 1969. Centenarian box tur
tles. Int. Turtle and Tortoise Society J. 3(3): 25-29.
Latham, R. and F.C. Schlauch. 1969. Inscribed eastern box tur
tles. Int. Turt. Tort. Soc. J. 3(4): 13.
Legler, J.M. 1960. Natural history of the ornate box turtle, Ter
rapene omata omata Agassiz. Univ. Kansas Publ. Mus. Nat.
Hist. 11: 527-669.
Metcalf, E. and A.L. Metcalf. 1970. Observations on ornate box
turtles (Terrapene omata omata Agassiz). Kansas Acad.
Sci. Trans. 73(1): 96-117.
Milstead, W.W. 1967. Fossil box turtles (Terrapene) from cen
tral North America, and box turtles of eastern Mexico.
Copeia 1967: 168-179.
__________ 1969. Studies on the evolution of box turtles (genus
Terrapene). Bull. Fla. State Mus., Biol. Sci. 14(1): 1-113.
__________and D.W. Tinkle. 1967. Terrapene of western Mexico,
with comments on the species groups in the genus. Copeia
1967: 180-187.
Nichols, J.T. 1939a. Data on size, growth and age in the box tur
tle, Terrapene Carolina. Copeia 1939: 14-20.
. 1939b. Range and homing of individual box turtles.
Copeia 1939: 125-127.
Oliver, J.A. 1953. Young Billy Johnson’s old box turtle. Animal
Kingdom 56(5): 154-156.
238
__________. 1955. The natural history of North American am

phibians and reptiles. D. van Nostrand Co., Princeton, N.J.
359 pp.
Webb, R.G., W.L. Minkley, and J.E. Craddock. 1963. Remarks
on the Coahuilan box turtle, Terrapene coahuila (Testu-
dines, Emydidae). Southw. Naturalist, 12(2): 89-99.
RHINOCLEMYS:
Alvarez del Toro, M. 1960. Los reptiles de Chiapas. Inst. Zool.
del Estado, Tuxtla Gutierrez, Chiapas, Mexico. 204 pp.
Boulenger, G.A. 1889. Catalogue of the chelonians, rhyncho-
cephalians and crocodiles in the British Museum (Natural
History). 311 pp.
Gray, J.E. 1861. On a new species of water-tortoise (Geoclem■
mys melanostema) from Darien. Proc. Zool. Soc. London
1861: 204-205.
__________. 1870. Notes on the species of Rhinoclemmys in the
British Museum. Proc. Zool. Soc. London 1863: 254-255.
McDowell, S.B. 1964. Partition of the genus Clemmys and rela
ted problems in the taxonomy of the aquatic Testudinidae.
Proc. Zool. Soc. London, 143: 239-279.
Medem, F. 1962. La distribucion geographica y ecologia de los
Rev. Acad. Colombiana Cien. Exactas, Fis. y Natur. 11(44):
279-303.
Mertens, R. 1954. Zur Kenntnis der Schildkrotenfauna Venezu-
elas. Senck. biol. 35(1/2): 3-7.
Moll, E.O. and J.M. Legler. 1971. The life history of a
neotropical slider turtle, Pseudemys scripta (Schoepff), in
Panama. Bull. Los Angeles Co. Mus. Nat. Hist. Sci., No. 11:
1-102.
Pritchard, P.C.H. 1964. Turtles of British Guiana. J. British
Guiana Mus. and Zoo, No. 39: 19-32.
Schmidt, K.P. 1946. Turtles collected by the Smithsonian Bio
logical Survey of the Panama Canal Zone. Smithsonian
Misc. Coll. 89(1): 1-20. Washington.
EMYS:
Bojanus, L.H. 1970. Anatome Testudinis Europaeae. Reprint
Edition, Society for the Study of Amphibians and Reptiles,
Ohio. Pp. 1-178.
cephalians, and crocodiles in the British Museum (Natural
History). 311 pp.
239
www.Ebook777.com
Plastral view of Missouri slider, Pseudemys floridana hoyi.
OPPOSITE: Above: Adult female Florida cooter, Pseudemys floridana penin-

Below: Adult Missouri slider turtle, Pseudemys floridana hoyi.
sularis.
240
www.Ebook777.com
Chaignon, H. de. 1904. Contributions a l’Histoire naturelle de la
Tunisie. Bull. Soc. Hist. Nat. Autun, 17: 1-166.
Gadow, H. 1901. Amphibia and Reptiles. In The Cambridge
Natural History. London, 8, xiii + 668 pp.
Hellmich, W. 1962. Reptiles and amphibians of Europe. Bland-
ford Press, London. 160 pp.
Loveridge, A. and E.E. Williams. 1957. Revision of the African
tortoises and turtles of the suborder Cryptodira. Bull. Mus.
Comp. Zool., 115(6): 163-557.
Pritchard, P.C.H. 1966. Notes on Persian Turtles. Brit. J.
Herpetology, 3(11): 271-275.
Rollinat, R. 1934. La vie des reptiles de la France Central.
Paris.
Schreiber, E. 1912. Herpetologia europaea. Jena: Gustav
Fischer; 946 pp.
MAUREMYS:
Anderson, S.C. 1963. Amphibians and reptiles from Iran. Proc.
Calif. Acad. Sci., 4th. Ser., XXXI (16): 417-498.
Gadow, H. 1901. op. cit.
Hellmich, W. 1962. op. cit.
Mao, S.H. 1971. Turtles of Taiwan. Commercial Press, Taipei;
128 pp.
McDowell, S.B. 1964. Partition of the genus Clemmys and rela
ted problems in the taxonomy of the aquatic Testudinidae.
Proc. Zool. Soc. London., 143: 239-279.
Nakamura, K. 1934. On Clemmys mutica (Cantor) with special
reference to its variations. Annot. Zool. Japan, 14: 425-435.
tral Asia. Amer. Mus. Nat. Hist., 10: 604 pp.
Pritchard, P.C.H. 1966. op. cit.
Schreiber, E. 1912. op. cit.
Smith, M.A. 1931. The Fauna of British India, including Ceylon
and Burma. Reptilia and Amphibia. Vol. I, Loricata, Testu-
dines. London: Taylor and Francis. 185 pp.
Stejneger, L. 1907. Herpetology of Japan and adjacent terri
tory. Bull. U.S. Nat. Mus., 58: 577 pp.
Villiers, A. 1958. Tortues et crocodiles de l’Afrique Noire Fran-
caise. I.F.A.N., Dakar. 354 pp.
SAC ALIA:
McDowell, S.B. 1964. op. cit.
Pope, C.H. 1935. op. cit.
ANNAMEMYS:
Bourret, R. 1939. Notes herpetologiques sur l’lndochine Fran-
caise, XVIII. Reptiles et batraciens recus au Laboratoire
des Sciences Naturelles de l’Universite au cours de l’annee
1939. Descriptions de quatre especes et d’une variete nouvel-
242
les. Bull. Gen. Instr. Publ., 1939(4): 5-39.

__________ 1941. Les Tortues de l’lndochine. Inst. Oceanogr.
l’lndochine, Note 38, 1-235.
Savage, J.M. 1953. Remarks on the Indo-Chinese turtle Anna-
memys annamensis; with special reference to the status of
Cyclemys annamensis. Ann. Mag. Nat. Hist., Ser. 12, vi:
468-472.
Siebenrock, F. 1903. Schildkroten des ostlichen Hinterindien.
Sitz. Akad. Wiss. Wien. Mathem-naturw. Klasse, 112(1):
4, 333-352.
HARDELLA:
Minton, S. 1966. A contribution to the herpetology of West Paki
stan. Bull. Amer. Mus. Nat. Hist. 134(2): 27-184.
Smith, M.A. 1931. op. cit.
BATAGUR:
Anderson, J. 1878-9. Anatomical and zoological researches and
zoological results of the Yunnan Expeditions. Reptilia and
Amphibia, pp. 705-860. Calcutta.
Hendrickson, J.R. 1961. Conservation investigations on Malay
an turtles. Malayan Nature Journal, Special Issue. Pp. 214-
223.
Maxwell, F.D. 1911. Reports on inland and sea fisheries in the
Thongwa, Myaungmya, and Bassein Districts and Turtle-
Banks of the Irrawaddy Division. 57 pp. Rangoon, Govern
ment Printing Office.
CALLAGUR:
Rooij, N. de. 1915-17. The reptiles of the Indo-Australian Archi
pelago. Vol. 1. Lacertilia, Chelonia, Emydosauria. 1915. Pp.
I-XIV, 1-384.
ORLITIA:
Rooij, N. de. 1915-17. op. cit.
Mehrtens, J.M. 1970. Orlitia the Bornean Terrapin. Int. Turt.
Tort. Soc. J. 4(5): 6-7,33.
KACHUGA:
Mertens, R. 1969. Eine neue Rasse der Dachschildkrote, Kachu-
ga tecta. Senckenbergiana biol. 50 (1/2): 23-30.
Minton, S. 1966. op. cit.
Spoczynska, J.O.I. 1969. The fascinating Kachugas. Int. Turt.
Tort. Soc. J. 3(2): 8-11.
Wermuth, H. 1956. Die Sud-Rasse der Indischen Dachschild
krote. Aquar. Terrar., Leipzig & Jena, 3: Nr. 8.
GEOEMYDA, HEOSEMYS:
McDowell, S.B. 1964. op. cit.
243
www.Ebook777.com
Two views of the river cooter Pseudemys concinna mobiliensis.
Young red-bellied turtle,

Pseudemys rubriventris,
from the northeastern
United States.
Below: Hatchling Florida

red-bellied turtle, Pseu-
demys nelsoni, from
Juniper Springs, Florida.
www.Ebook777.com
Taylor, E.H. 1920. Philippine turtles. Philippine J. Sci., 15:
1-144.
__________ 1970. The turtles and crocodilians of Thailand and
adjacent waters. Univ. Kansas Sci. Bull. XLIX: 87-179.
MELANOCHELYS:
Deraniyagala, P.E.P. 1939. The tetrapod reptiles of Ceylon.
Colombo, 412 pp.
CUORA, CYCLEMYS:
Mao, S.H. 1971. op. cit.
Taylor, E.H. 1920. op. cit.
__________ 1970. op. cit.
HIEREMYS, NOTOCHELYS:
OCADIA:
Mao, S.H. 1971. op. cit.
MORENIA:
Anderson, J. 1879. op. cit.
Klingelhoffer, W. 1959. Terrarienkunde, 4 Teil; Stuttgart.
GEOCLEMYS:
Minton, S.A. 1966. op. cit.
CHINE MYS:
Mertens, R. 1936. Riesenwuchs bei einer Schildkrote, Chinemys
reevesii Gray. Zool. Anz., Leipzig 114(7/8): 206-209.
Priemel, K. 1918. Fortpflanzung von Geoclemys reevesi in
Gefangenschaft. Wschr. Aquar. Terrar.-Kunde 15: 157-158.
MALAYEMYS, SIEBENROCKIELLA:
246
Chapter 5
LAND TORTOISES
The land tortoises, which comprise the family Testudinidae,
are the only turtles fully adapted for a terrestrial existence, except
for a few species of the Emydidae. Several distinctive features of
this family may be mentioned: the hard, horny shells (frequently
with well-marked growth annuli on each scute); the predominant
ly or exclusively herbivorous diet; the almost exclusively terres
trial habitat ; and the characteristic shape of the feet, the front legs
slightly flattened from front to back and with very thick heavy
scales on the anterior surface. The hind limbs are normally colum
nar and elephantine in shape and have loose skin which folds con
certina-fashion when the limbs are retracted. When a tortoise has
completely retracted its head and limbs, the only exposed parts
are the soles of the hind feet, the tail region, and the scaly anterior
surfaces of the forelimbs, which are drawn together at the elbows,
completely covering the head. Some tortoises have improved even
on this protection, Kinixys by having a hinge across the carapace
so that the rear part can be lowered to protect the hind limbs,
Pyxis by having a moveable hind plastral lobe, and Geochelone
denticulata, carbonaria, pardalis, and some others by having a
thickened and raised gular lip and downcurved pygal region. The
tortoise thus depends upon its armor rather than on an aggressive
or flight reaction for protection, and it is about the only vertebrate
that one can walk up to in the wild and pick up without difficulty.
This ease of capture, together with convenience of storage and
tasty flesh, has had the result that the larger tortoises of many
tropical areas are becoming rarer each year and in many cases
have disappeared completely from the vicinity of human habita
tions. Temperate zone tortoises too are heavily exploited for the
pet trade; tortoises are popular pets because of their quaint and
harmless disposition, but most species are much harder to main
tain in captivity for prolonged periods than their aquatic counter
parts, unless they can be kept in an area within their natural range
or which provides a reasonable facsimile of the climate in their
natural habitat. Correct humidity, temperature, and sunlight are
all vital for maintenance of the health of captive tortoises.
Many authors ‘lump’ the tortoises in the same family as the
emydid turtles, arguing that the structural differences between
247
www.Ebook777.com
Midland painted turtle, Chrysemys picta marginata.
Alabama red-bellied turtle, Pseudemys alabamensis, a species probably con

fined to the vicinity of Mobile Bay. Photo by Dr. R.W. Barbour.
248
Adult female Barbour’s map turtle, Graptemys barbouri, from the Chipola
River, Florida.
Adult female (left) and adult male Alabama map turtle, Graptemys pulchra.
www.Ebook777.com
the two groups are minimal. However, one structural difference
always separates the two groups: the number of phalanges in the
digits of the hind foot is never more than two in the tortoises, but in
the Emydidae one toe at least always has more than two phalan
ges. This trivial but constant difference, together with their radi
cally different physiology, justifies separation of the two families.
It is usually assumed that tortoises are derived from emydid
turtles. It seems surprising that such an ancient and completely
terrestrial group, derived originally from primitive reptiles which
themselves were terrestrial, should have had an aquatic phase in
its phylogeny, but such appears to be the case. The family is
thought to have originated in Southeast Asia, where some of the
most primitive living tortoises (e.g. Geochelone emys) are still
found. Certain tortoise species are considered ‘primitive’ in that
they show certain characteristics typical of generalized emydids.
Loveridge and Williams list the following criteria for primitive
ness in tortoises:
OSTEOLOGICAL CHARACTERS
i) Interval between ventral processes of the prefrontals only
moderately widened.
ii) Maxilla with triturating surface ridged, but premaxilla un
ridged.
iii) Anterior palatine foramina small, concealed.
iv) Temporal arcade strong.
v) Prootic well exposed dorsally.
vi) Quadrate not enclosing stapes.
vii) Surangular subequal in height to prearticular.
viii) Fourth cervical centrum biconvex.
ix) Anterior neurals hexagonal.
x) Suprapygal single, anterior to the vertebral-supracaudal
sulcus.
xi) Entoplastron anterior to humeropectoral sulcus.
xii) Digital phalangeal formula 2,3,3,3,3.
EXTERNAL CHARACTERS
i) At least prefrontal and frontal head scales present.
ii) Scales on forelimb numerous, not greatly enlarged.
iii) Large femoral tubercles present.
iv) Claws on forelimb 5, on hind limb 4.
v) Tail claw absent.
vi) Carapace moderately convex.
vii) Neither carapace nor plastron hinged.
viii) Nuchal present.
ix) Vertebrals not conical.
x) Vertebral keel very weak, lateral keels absent.
xi) Supracaudal divided.
250
At the present time, ten genera of living tortoises are recog

nized. The family reaches its greatest diversity in Africa, where
there are five endemic genera (Kinixys, Malacochersus, Cher sine,
Homopus, and Psammobates). Madagascar has two more en
demic genera (Pyxis and Acinixys), and North America one
(Gopherus). The genus Testudo is found on both European and
North African sides of the Mediterranean and extends as far east
as Pakistan. Geochelone is the largest and most widely distributed
genus, with two species in Africa, two in Madagascar, one (and
various extinct forms) in the islands of the Indian Ocean, six in
Southeast Asia, one in Celebes, three in South America, and one
polytypic species in the Galapagos Islands. All the larger tortoises
belong to this genus, and their large size may have been contribu
tory to the extinction of the genus in such areas as North America,
where present-day winters are cold; the surviving North Ameri-
251
www.Ebook777.com
Young of three Graptemys species. Above: Graptemys pulchra, the Alabama
map turtle; the height of the dorsal ridge varies from one river system to
another. Opposite, above: Graptemys oculifera from the Pearl River on the
Louisiana-Mississippi border. Opposite, below: Graptemys flavimaculata, the
yellow-blotched sawback.
252
www.Ebook777.com
can tortoises, all belonging to the genus Gopherus, have specially
adapted forefeet which enable them to burrow to escape cold
nights—or cold winters.
The occurrence of gigantic tortoises-and only gigantic ones-
on oceanic islands is a phenomenon worth further discussion.
Before the islands were despoiled by man in the 18th and 19th cen
turies, giant tortoises were found on dozens of islands in the wes
tern Indian Ocean, and they are still found on most of the larger
Galapagos Islands, in the east Pacific Ocean. There are a number
of criteria which apparently have to be fulfilled for an island to
have giant tortoises: i) the island must be in the tropics, since
giant tortoises have no means of escaping very cold weather;
ii) the island must be several hundred miles from the nearest
mainland; if the island is too close to the mainland, it will have a
continental-type fauna, including perhaps carnivorous mammals,
whose presence is apparently inimical to giant tortoise survival;
on the other hand, islands too far from the nearest mainland (such
as Ascension and Hawaii) are apparently beyond the reach of the
powers of dispersal of tortoises; iii) there must have been a popu
lation of large tortoises on the nearest mainland area; Madagas
car, which had truly gigantic tortoises until comparatively recent
ly, may have been originally colonized from Africa, and later
acted as a secondary dispersal center for the Mascarenes,
Aldabra, the Seychelles, and the other formerly tortoise-bearing
islands of the Indian Ocean.
Whether or not the tortoises were already gigantic when they
first reached the islands cannot be stated with certainty, since the
geological conditions of both the Galapagos and the Indian Ocean
Islands render the preservation of fossils extremely unlikely. How
ever, the best educated guess one can make is that the tortoises
were already very large when they arrived. For one thing it
should be borne in mind that in prehistoric times tortoises fully’ as
large as those of Galapagos or Aldabra lived in many parts of the
world. Geochelone crassiscutata, for example, was a four-foot long
tortoise from the Pleistocene of Florida and Texas; Geochelone
cubensis was a giant tortoise from the Pleistocene of Cuba- Geo
chelone ammon was a giant tortoise from Egypt; G. perpiniana
another giant species from the south of France; while the largest
of all, G. sivalensis (or Colossochelys atlas) was a six-foot tortoise
from the Pliocene of northern India. Even today, the distinction
between island ‘giant’ tortoises and mainland ‘small’ tortoises is
not as clear-cut as many people believe. For example, some Gala
pagos tortoises are certainly huge; on Albemarle Island males
may reach a straight carapace length of over fifty inches; on the
other hand, on Hood Island even males do not grow longer than
about thirty inches, while females are mature at about 21 inches.
254
By comparison, the tortoises in adjacent Peru on the mainland of

South America may on occasion reach a carapace length of about
thirty inches, and in extreme cases reach a weight of well over 100
lbs. (though on the mainland, interestingly, it is the female that is
by far the larger). In Africa too the spurred tortoise (Geochelone
sulcata) may exceed a carapace length of thirty inches.
Another reason for believing that gigantism was a pre-adapta
tion for island life rather than a result of it is that a large tortoise is
much more likely to survive passive flotation across several hun
dred miles of ocean than a small one; it can hold greater food re
serves, can raise its head higher above the waves to avoid drown
ing, and so forth. The theory of passive flotation to the Galapagos
and the Indian Ocean Islands has been discredited by some, but
the prevailing current patterns support such a theory, and even
today tortoises are quite often seen taking voluntarily to the sea in
the main lagoon of Aldabra Island. Once having arrived on an
oceanic island, moreover, a giant tortoise would find its size a use
ful preadaptation both to enable it to cross the extremely fissured
and precipitous terrain of both volcanic and coral oceanic islands,
and to help it damp out effects of the often marked day-night temp
erature changes of equatorial islands; it is doubtful if any pres
ently-existing small mainland tortoise could long survive the cool,
all-pervading dampness of the highlands of certain Galapagos
Islands. A large tortoise is also at an advantage if it should have to
traverse an extensive shadeless piece of terrain, since it would
heat up more slowly in the sun. The Galapagos and Indian Ocean
Islands are (or were) the only ecosystems in the world in which
tortoises are the dominant herbivores; the newly-arrived ances
tral Galapagos or Aldabra tortoise found abundant food, few com
petitors, and even fewer predators, and in such a simple eco
system they soon reached almost incredible population densities.
Even in the Galapagos, where most if not all of the populations are
markedly reduced by the activities of man and his feral animals, it
is still possible to see more tortoises in a single day than in almost
any mainland area of equal extent; the exceedingly high popula
tion density of tortoises on Aldabra at the present time is discussed
later in this chapter.
Tortoises are in fact better at colonizing islands than any fami
ly of freshwater turtles. This may be in part because the fresh
water niche is a restricted or absent one on many small islands,
but it may also be because a freshwater turtle which, by some
accident, finds itself at sea, may simply ‘feel at home’ in this
watery medium, and start behaving as it normally would in
water—that is, diving to the bottom and interacting osmotically
with its environment, with fatal results. However, a few emydid
turtles (Malaclemys, Callagur, Pseudemys concinna suwannien-
255
www.Ebook777.com
An adult male black-knobbed sawback turtle, Graptemys nigrinoda, from
Alabama.
256
Juvenile common map turtle, Graptemys geographies , from the north-central

United States.
Adult female common map turtle, Graptemys geographies.
257
www.Ebook777.com
sis) take to brackish or salt water more or less regularly, and one
trionychid (Pelochelys bibroni) is so at home at sea that it has suc
ceeded in penetrating from the Malay Peninsula through the Indo
nesian islands as far as New Guinea, while Trionyx triunguis is
regularly caught in deep water in the Mediterranean.
Many tortoises show an interesting modification of the neural
bones which, instead of being hexagonal throughout the series as
in emydids, become alternately quadrilateral and octagonal, at
least toward the middle of the series. The pleural bones corres
pond to this alternation, the even-numbered ones having broad
distal ends and narrow proximal ends which suture with the out-
ward-facing sides of the octagonal neurals, while the odd-number
ed ones have narrow distal ends and broad proximal ends, which
suture with the outward-facing sides of the quadrilateral neurals
and also with the adjacent sides of each of two octagonal neurals.
This neural modification is found in all living tortoise genera
except Kinixys, Acinixys, Homopus, and Psammobates; sugges
tions of it are even present in the bizarrely-modified Malacocher-
sus. In Gopherus berlandieri the alternation in shape of the
pleurals may be so extreme that the odd-numbered pleurals with
narrow distal ends become closed out completely from any contact
with the peripheral bones by the greatly broadened distal ends of
the even-numbered pleurals. On the other hand, the octagonal-
quadrilateral arrangement is often imperfectly or asymmetrically
expressed; for example, a carapace of Geochelone elephantopus
microphyes in my possession does not have a single neural bone
that is either octagonal or quadrilateral, the normal pattern being
distorted into one of asymmetrical pentagons and heptagons. The
precise adaptive significance of octagonal/quadrilateral neurals
is not obvious, but presumably it helps stabilize the carapace
against transverse cracking.
GENUS GEOCHELONE
The genus Geochelone is defined by a complex of characters;
these include the strongly ridged triturating surface of the maxil
la, the maxilla not entering the roof of the palate, the small an
terior palatine foramen concealed in ventral view, the prootic typi
cally well exposed dorsally and anteriorly, the quadrate usually
enclosing the stapes, the surangular subequal in height to the pre-
articular, the biconvex 2nd, 3rd, and 4th cervical vertebrae, the
unhinged carapace and plastron, the typically alternating octago
nal/quadrilateral pattern of the anterior neurals, the outer edge of
the third costal about the same length as, or longer than, that of the
fourth costal, the two suprapygals, with the anterior one larger
than the posterior, which it partially encloses between its postero-
laterally directed arms, the sulcus between the fifth vertebral and
the supracaudal opening near the middle of the second supra-
258
pygal, the thickened produced gular region, and the single or

double gular scutes which are longer than broad.
Six living and one subfossil subgenera are included within the
genus Geochelone: the subgenus Geochelone itself, from tropical
Africa and the Indian region; Asterochelys, from Madagascar;
Aldabrachelys, from certain Indian Ocean Islands; Cylindraspis,
formerly of the Mascarenes; Manouria and Indotestudo, both from
southeast Asia; and Chelonoidis, from South America and the
Galapagos Islands.
SUBGENUS GEOCHELONE
The subgenus Geochelone is defined by the following combina
tion of features: alternately octagonal and quadrilateral anterior
neurals; entoplastron not crossed by humeropectoral sulcus;
supracaudal undivided; gulars paired but not divergent; first dor
sal vertebra short and stout; nuchal absent; external naris not
higher than wide; quadrate enclosing stapes; postotic fenestra not
concealed in posterior view. Four species are included—two in
Africa and two in Asia. The African species are much larger than
the Asiatic and lack a claw on the tip of the tail.
The African species are Geochelone sulcata, the spurred tor
toise, and Geochelone pardalis, the leopard tortoise. The spurred
tortoise is the largest mainland tortoise; adult specimens are com-
The African spurred tortoise, Geochelone sulcata. Geochelone calcarata is a

synonym.
www.Ebook777.com
Two subspecies of the false map turtle from the central United States. Above:
Graptemys pseudogeographica ouachitensis. Below: Graptemys
pseudogeographica pseudogeographica.
260
www.Ebook777.com
monly well over 24 inches in length, and Stanley Flower reports an
individual that, when nearly eighteen years old, had a straight
carapace length of exactly thirty inches and weighed 184 lbs. The
largest specimens are bigger still, but precise measurements are
lacking. This species has a wide, rather flattened carapace of a
uniform, unmarked yellow or horn color. In fully mature speci
mens, males especially, the anterior marginal scutes become
strongly reverted. Growth annuli on the scutes of the carapace are
usually very well defined. In the plastron, the gulars are strongly
forked, the anals have a deep median notch, and the pectoral
scutes are extremely narrow. The head is moderately large and
the upper jaw is doubly hooked. The head and limbs are the same
color as the carapace and are well protected with thick scales. The
species derives its name from the presence of two or three strong
spurs on each side of the tail. It is an arid country species, as its
sandy coloration would suggest, and is found right across the
southern fringes of the Sahara, from inland Senegal through Maur
itania, Mali, Niger, Chad, the Sudan, and Ethiopia to the Massaua
coast on the Red Sea. Since much of the country where this tortoise
is found is inhabited by Moslems, it is possible that it is not as
heavily exploited for food as are other large tortoises. There are
large specimens in a number of the bigger zoos in Europe, but very
few are ever imported into the United States. One individual sur
vived 42 years in captivity. This species is maintained in large
numbers at the Bamako Zoo (Mali), where reproduction frequent
ly occurs in the very large open-air enclosures. When Karl Switak
visited this zoo in 1969, he was shown forty of fifty young spurred
tortoises that had hatched in the zoo and which would soon be re
leased in their native environment.
The habits of the spurred tortoise in the wild are not very well
known. Much of the habitat is so dry that it may not have access to
standing water for years at a time. It avoids excessive moisture
loss by having a highly impermeable skin and also by digging ‘pal
lets’ or shallow burrows in the ground. A typical adult pallet slopes
down to a maximum depth of about thirty inches, though Flower
mentions the figure of four yards for one pallet and the eleventh
century writer El Bekri (quoted by Villiers) mentions burrows so
huge that a man could walk about inside them and tortoises so big
that the combined efforts of several men, pulling on ropes, were
necessary to remove them! The pallets are excavated with the
front limbs, presumably assisted by the strongly projecting and
forked gulars.
Copulation takes place during the rainy season and may be
repeated several times in the course of one day, each contact
taking about fifteen minutes. As with the giant tortoises of oceanic
islands, the male is much larger than the female, and has a deep
262
plastral concavity. During copulation the female is virtually im

mobilized by the weight of the male, who presses down on the an
terior part of the female’s carapace with his forelimbs. While in
this position, the male utters rhythmic low-pitched sounds that
have been described as muffled rattling groans. The egg-laying
process has not been described, but the hatchlings have pale yel
low carapaces, each scute with narrow brown borders. The shell of
the hatchling is almost two inches in length and only slightly nar
rower than long; the entire periphery of the carapace is wavy or
serrated.
In captivity the spurred tortoise eats a wide variety of vegeta
tion, preferring (according to Villiers) the leaves of the morning-
glory (lpomaea)\ it will also drink water more frequently and in
greater quantities than it would ever have access to in the wild.
The other African member of the subgenus Geochelone is the
leopard tortoise, Geochelone pardalis. It does not reach quite as
large a size as the spurred tortoise, but nevertheless it is a good-
sized animal, the largest adults being about two feet long and
weighing about seventy pounds. Nevertheless, specimens found in
the wild are usually much smaller than this, and one author re
ports that some individuals remain at about ten pounds, though
others grow much larger. On the other hand, W.H. Archer has
recently described a population of leopard tortoises in the Graaff-
Reinet area of South Africa which includes some truly gigantic
specimens; one measured 26 inches and weighed 96 lbs., while ano
ther weighed 105 lbs. (African Wild Life, Sept. 1968, Vol. 22, No. 3).
The sexes are not obviously different in adult size. Numerous char
acteristics distinguish the sexes in the majority of cases, but the
263
www.Ebook777.com
Cagle’s map tur
tle, Graptemys
caglei, a newly
described species
from Texas.
Plastral views of diamondback terrapins, Malaclemys terrapin, showing varia

tion in shell and skin pattern.
Diamondback terrapin, Malaclemys terrapin macrospilota, from Tampa Bay,

Florida. This subspecies is distinguished by the light areolae of the carapace
scutes, but the white head is only shown by certain individuals.
Heads of four ornate diamondback terrapins, Malaclemys terrapin tequesta,

all from Merritt Island, Florida. Photo by Dr. Llewelyn Ehrhart.
www.Ebook777.com
slightly concave posterior third of the plastron and the longer tail
of the male are the only ones that hold for all cases.
The leopard tortoise is distinguished from G. sulcata by the
much more domed carapace, which never acquires reverted mar
gins, and by the presence of black pigmentation on the carapace.
The ground color of the carapace is tawny-yellow or buff, while the
areolae are usually brown. The black markings consist of more or
less unbroken lines around the carapace scutes in young speci
mens, but in older and adult tortoises they consist of irregular bro
ken radiations or spots. They may become so dense as to almost
obscure the ground color; on the other hand Ethiopian leopard tor
toises tend to have very sparse shell pigmentation. In the Graaff-
Reinet area of South Africa, the tortoises found at altitudes of over
6000 feet have much more distinct markings and reach a much lar
ger size than those found below 4000 feet.
The leopard and the spurred tortoises are nowhere sympatric,
though their ranges approach each other in Ethiopia. The range of
the leopard tortoise extends over much of south and east Africa,
having been recorded from the southern parts of the Sudan and
Ethiopia, the Somali Republic, Uganda, Tanzania, Kenya, Mozam-
Large leopard tortoise, Geochelone pardalis babcocki, with the domed

carapace typical of the subspecies.
266
bique, Malawi, Zambia, Rhodesia, Botswana, Natal, Transvaal,

Cape Province, southwest Africa, and Angola. The typical habitat
is savanna, with more vegetation than is typical of the range of sul
cata. Two subspecies are currently recognized; the typical form,
Geochelone pardalis pardalis, is a rarely-seen animal from south
west Africa north to around Kolmanskop and Keetmanshoop. Geo
chelone pardalis babcocki occupies the remainder of the range of
the species; doubtless an intensive study of variation in the leop
ard tortoise would lead to the proposal of a number of new names.
G.p. babcocki is distinguished by the more domed carapace of the
adult and by the carapace pattern of the young in which the dark
spots on the central scutes are located along the midline instead of
on either side of it.
Juvenile leopard tor

toise, Geochelone
pardalis pardalis.
Notice the spot pat
tern. Photo by John
Visser.
Karl Switak records that the diet of the leopard tortoise in

Kruger National Park, South Africa, includes 'the Cotyledons and
other succulent plants, but (they) have been known to feed on toad
stools and the fallen fruit of Sclerocarya birrea trees.’ Hewitt
found that leopard tortoises feed principally on grass, but also on
various succulents, crassulas, spekboom, thistles, prickly pear,
pumpkins, beans, and watermelons. Some captive specimens eat
smashed chicken bones, and those specimens which are regularly
supplied with a bone supplement in their diet may grow very
rapidly.
During courtship, males compete for females by pushing and
butting each other until one is overturned. One observer has also
reported that large females may butt and ram other tortoises of
both sexes. During copulation itself, the neck is extended low over
the carapace of the female and the husky cry uttered is audible
some distance away. Nesting takes place in spring (September
and October) in South Africa, but the season is much longer in
267
www.Ebook777.com
'
Adult female chicken turtle, Deirochelys reticularia, showing the vertical

striping on the rump, a useful identification character.
Deirochelys reticularia chrysea, the Florida chicken turtle.
268
The American wood turtle, Clemmys insculpta, a terrestrial species from the
northeastern United States.
Blanding’s turtle, Emydoidea blandingi, a semi-aquatic species from the

Great Lakes region.
269
www.Ebook777.com
tropical Africa. As with other tortoises, preparation of the nest is a
protracted and seemingly very inefficient business; the female
urinates on the soil to facilitate excavation, but the elephantine
shape of the hind limbs is far from ideal for digging, and the
amount of soil withdrawn by the tortoise during each excavatory
movement is minimal. Two hours or more may elapse before the
cavity is ready. The eggs are almost spherical, very hard-shelled,
and average about 1.4” in diameter. Recorded clutches have
ranged in number from five to thirty, while one captive tortoise
laid a total of 52 eggs in six installments during one season. When
the number of eggs in a clutch is very high, they may be deposited
in two or more discrete layers separated by earth. Archer reports
that two female leopard tortoises of comparable size but from dif
ferent localities (Transvaal and Eastern Cape respectively) pro
duced eggs of very different size and number; the former always
laid between 12 and 24 small eggs, 1” to 1 1/8” in diameter, while
the latter laid 8 to 12 eggs, each 1.75” to 2” in diameter.
From eight to as many as eighteen months may elapse before
the hatchling tortoises emerge at the surface. Hatchlings and juve
niles fall prey to ground hawks and crows in South Africa, while
the principal enemies of the adults are ratels, hyaenas, bush fires,
and, of course, man. Human predation is probably responsible for
the relative rarity of the leopard tortoise in many easily-accessi-
ble, low-lying areas.
The leopard tortoise has survived for 75 years in captivity.
This species has a well-developed homing instinct, and Archer re
ports that several individuals translocated five to eight miles away
were back at the point of origin within a fortnight, even though to
get back they had to climb over a 4 ft. high fence of 3” mesh. Such
fences are climbed with little trouble, the animal simply hooking
its feet into the mesh as it goes up, then toppling over once it reach
es the top; even if it lands on its back, it is able to right itself with
out much trouble.
The Indian star tortoise, Geochelone elegans, is closely related
to the leopard tortoise, but the superficial differences render the
species immediately distinguishable. The star tortoise is found in
Ceylon and peninsular India, extending west as far as Sind. It is
sometimes very common on small offshore islands, such as Kara-
duva and Ramaswaran. It is a comparatively small species; a
very large female may be ten inches long, a large male about six
inches. This is the most familiar of the so-called ‘star’ tortoises, so
named because of the pattern of the shell. Each of the large scutes
of the carapace has a yellow areola from which a number of radia
ting yellow streaks extend to the margins of the scute; the back
ground color of the carapace is black or dark brown. Subsidiary
radiating streaks may extend from intermediate points to the
270
scute margins; whenever a discontinuity appears in the dark pig

mentation at the edge of a scute, this discontinuity is propagated
as a radial streak as the scute grows. The radiated pattern is also
present on the marginal scutes, though there the areolae are situa
ted at the postero-ventral comers, and the ‘stars’ are incomplete.
The radiations are continued on the plastron. The forelimbs bear
large scales and are yellow in color. The head, which is somewhat
bulging in the nasal region, is basically yellow, but there may be
black spots on the crown. The females are not only much larger
than the males, but also have broader shells, and frequently the
vertebral scutes and to a lesser extent the costals, are conical in
shape, the areola forming the vertex in each case. Males have nar
row, relatively smooth shells and often have rather dingy mark
ings compared to those of the females. In males the supracaudal
scute is extended downward over the tail.
The shell pattern of the star tortoise is very striking when the
animal is in hand, but in the wild the yellow radiations provide ef
fective camouflage when the tortoise is partially concealed under
tussocks of dry grass. As with many arid-land tortoises, feeding
and active moving about take place in the early morning and late
The Indian star tortoise, Geochelone elegans, once one of the most familiar
exotic tortoises in captivity.
271
www.Ebook777.com
The spotted turtle, Clemmys guttata , a familiar, diminutive species of the
eastern United States.
272
The California pond turtle, Clemmys marmorata, one of the few freshwater
turtles from the western United States. Photo below by Ken Lucas, Steinhart
Aquarium.
273
www.Ebook777.com
afternoon hours, except during rain, when the tortoises may be
seen abroad at midday. According to Deraniyagala, the principal
diet is grass, fallen fruit or flowers, and such types of vegetation as
are easily accessible.
Copulation is reported to take place throughout the four
months of the rainy season, which extends from mid-June to mid-
October. Males may attempt to turn each other over on their backs
at this time also. The nesting process appears to be more or less
similar to that of the leopard tortoise, even in the female urinating
on the earth to assist the excavation process and the maintenance
of a cavity. The eggs are reported to number from three to six, and
are slightly longer than wide; length is approximately 1.8” and
width approximately 1.4”. The eggshells are very hard and are
dotted with numerous widely-spaced pores. Two or three clutches
may be deposited in the course of a season. Hatchlings lack ‘star’
markings on the shell; Deraniyagala gives the following descrip
tion of the coloration of young elegans in Sri Lanka:
“In the young there are two distinct color variations, an
orange form with a yellow vertebral stripe, and the yellow
form without it. Dorsally there are about five or six black
neural blotches encroaching upon the areolae and five black
costal blotches on each side suggesting the former presence
of five pairs of costals. The first and last blotches point an-
Geochelone platynota, the rare Burmese star tortoise. Specimen in the

British Museum (Natural History).
274
teriorly and posteriorly respectively. The yellow neural

hand extends from the margin of the carapace to the areola
of the last vertebral scute but one, it is there interrupted and
sometimes continued along the last vertebral almost to the
supracaudal margin. Ventrally there are five pairs of black
blotches near sutures between plastral scutes. ”
Geochelone platynota is a poorly-known species closely
related to G. elegans; it is widely distributed in Upper Burma and
is found south as far as Moulmein. The adult size is similar to that
of G. elegans, but the shell is broader and the carapace scutes are
not humped or conical in shape. The radiating pattern of the cara
pace is simpler than that of elegans, there usually being six or
fewer light radiations on each scute (usually eight or more in
elegans). The pattern on the plastron is simplified even further;
the plastral scutes are basically yellow, with a single, broad dark
brown or black area on each. As with G. elegans, the tip of the tail
is equipped with a claw.
The eggs of platynota are similar in shape, and probably in
number, to those of elegans; they are said to be deposited at the
end of February. This species is very rarely seen alive outside of
its native country. Blyth reported that Burmans were so fond of
eating this tortoise that it was very difficult to obtain a living speci
men, though empty shells were used in numbers for baling oil out
of earthen vessels.
SUBGENUS ASTEROCHELYS
The two large tortoises of Madagascar, Geochelone radiata
and G. yniphora, are referred to a separate subgenus, Astero-
chelys, which is distinguished from Geochelone by the presence of
a nuchal scute and by the broader pectoral scutes. Geochelone
radiata, the radiated tortoise, is an exceedingly beautiful animal
whose range is now apparently restricted to the arid Antandroy
Territory of southern Madagascar, between the Mandrare River
on the east and the Menarandra Stream on the west—an area of
about 7,600 square miles. In its natural environment the radiated
tortoise is subjected to very low temperatures in winter, but
having no escape from frost, its range does not extend into the
mountains beyond the frost line. Like the African tortoises, the
radiated tortoise is seen most frequently during periods of rain.
This is a large species, big adults of both sexes being about fifteen
inches long, while Robert Baudy, who has collected a large series
of this species for captive breeding purposes, mentions one male
specimen weighing about 28 pounds. The carapace is very highly
domed, but the individual scutes are not conical. The head is yel
low except for a black area on top, and the limbs, which are cover
ed with rather small, non-imbricate scales, are entirely yellow.
The carapace scutes have yellow areolae with rather numerous
275
www.Ebook777.com
The southwestern subspecies of the ornate box turtle, Terrapene ornata
luteola.
OPPOSITE: Above: Terrapene nelsoni klauberi from Sonora, a rarely

photographed species. Below: The ornate box turtle, Terrapene ornata
ornata.
276
277
www.Ebook777.com
yellow streaks radiating to the edges. On the marginal scutes the
yellow radiations are mostly directed upward, but on the costals
most of the yellow pigment is concentrated in the triangular area
between the areola and the two lower corners of the scute, though
there is usually also a fan of finer yellow streaks extending to the
upper part of the scute. The area between the yellow streaks is
very dark brown or black. The plastral pattern is not dissimilar to
that of G. platynota, but some ‘fine structure’ of black radiations is
present in the yellow areas, and vice versa. Adult males have a
deep plastral concavity. In old adults the black pigmentation of the
carapace may spread to cover all but the areolae; one specimen,
imported to Indochina, in which this happened and which more
over had been deformed by years in captivity, was described as a
separate species (Testudo hypselonota) by Bourret in 1941. Ano
ther long-misidentified individual of this species was Tui Malila,
the famous Tonga tortoise, which was reputedly presented to the
then Queen of Tonga by Captain Cook in 1773 or 1777, and which
lived until May 19, 1966, having survived at least one forest fire as
well as a kicking by a horse which severely damaged the shell. For
the last years of its life it was completely blind and had to be fed by
hand. Until 1958, when James Oliver established its true identity,
this tortoise was thought to be an Indian Ocean giant tortoise. Its
remains are now preserved in the Auckland Museum.
The radiated tortoise owes its survival to the present day to the
gentle nature of the Antandroy people, who venerate the tortoises
which they call ‘Sokakes,’ and who will not molest them under any
circumstances; some individual tortoises living near certain vil
lages have been known to the villagers for many decades. Until
1925 the tortoises also obtained considerable protection from the
almost impenetrable Opuntia cactus thickets which they frequent
ed; these cactus stands had been planted by the Antandroys as a
defensive measure, but they were wiped out by an invasion of the
beetle Dactylopus coccus in 1925. Chinese in Tananarive nowadays
will pay the equivalent of fifty dollars for a radiated tortoise for
eating purposes (they are believed to have aphrodisiac proper
ties), and despite laws completely protecting the species, traffic in
tortoises from the south coast to the capital continues; some are
even exported to Reunion.
In captivity the radiated tortoise is very partial to melon, but
will eat most vegetable foods when melon is not available. The pre
ferred diet in the wild state has not been studied, but it is probable
that the radiated tortoise feeds on grass and any other green vege
tation and fruits that its arid habitat presents. When freshly
caught, this tortoise emits high-pitched cries, sometimes for as
long as an hour after capture; but after this initial outburst, they
may be kept in captivity for years without the sound being heard
278
Comparison of the plastra of Geochelone radiata (left) and Geochelone

yniphora (right).
again. Baudy also reports that fresh-caught radiated tortoises void

large volumes of reddish or dark brown liquid when picked up,
though this habit too is immediately lost in captive specimens. R.
Paulian reports that in the wild the females lay about twelve eggs
in September; Baudy’s specimens, transported to the northern
hemisphere, laid in March. The Gladys Porter Zoo in Brownsville,
Texas, succeeded in hatching an egg of the radiated tortoise in
early 1973, and William Zovickian has recently published some
data on captive breeding of radiata in Colorado. The three nestings
observed by Zovickian took place in July, August, and September;
clutches numbered 3, 4, and 6, respectively, and the eggs were
almost spherical, 36 to 42 mm in greatest diameter and 32 to 39 mm
in least diameter. The nests were flask-shaped and 6 to 8 inches
deep, with a maximum width of about six inches. The excavation
was carried out with the hind feet, one hind foot being used until it
tires, then the other. Excavation was assisted by frequent urina
tion by the female. Two of the eggs hatched after 145 days; the
hatchlings were 114 and 1 5/16 inches in carapace length. The
limbs of hatchling radiata are without dark pigment, as are those
of the adults. The carapace is black and whitish-buff rather than
black and yellow; the light coloration is arranged in a thick, web
like pattern on the black background.
279
www.Ebook777.com
An American box turtle, Terrapene Carolina, intermediate between the
subspecies Carolina and bauri.
Specimen of the Florida box turtle, Terrapene Carolina bauri, from near
Gainesville, Florida.
280
A Mexican box turtle, Terrapene Carolina mexicana, from Tamaulipas, Mex

ico. Photo by Patrick Burchfield.
Gulf Coast box turtle, Terrapene Carolina major.
www.Ebook777.com
The courtship of G. radiata in captivity was described by
Zovickian as follows:
“Once the specimens have reached sexual maturity,
sexual activity is begun. The males will attempt to mate
when their straight carapace length is about 12”; however
they are seldom successful until their length is 13” or more.
One reason for this is inexperience; another is that the fe
males do not mate until they approach 13.5” in length. The
smaller males are simply unable to effect matings with the
larger, stronger females.
“As in most testudines, the overtures of courtship are
performed by the male. He circles the female many times,
occasionally emitting a ‘clucking’noise. During the circling,
he often attempts to lift the female’s carapace with his gular
scutes and anterior marginals in an effort to keep her from
moving away. Actual mating occurs when the female re
sponds by raising herself up on all fours. This enables the
male to mount her and place his tail under hers. At this time,
the male alternately extends and suddenly relaxes his hind
legs causing his plastron to strike the female’s carapace
with a sharp thud. A very loud hiss is emitted during this
stage, so that the complete cycle is very reminiscent of the
sound made by a steam pile driver. Once intromission is ac
complished the male stops hissing and remains mounted,
often with mouth agape. The intromission lasts an average
of eight minutes, although entire mating behavior may take
over three hours. ”
Geochelone yniphora is somewhat similar in size and general
shape to G. radiata, but the highly domed carapace is slightly nar
rower and is uniform sandy brown in color, and the gular scute is
single, elongate, rounded anteriorly, and curved upward toward
the neck. This tortoise is confined to the Sakalav Territory in
northwest Madagascar and must now be considered one of the
rarest tortoises in the world. However, despite rumors to the con
trary, a small pocket of individuals still exists, and at the time of
writing three have been captured by the Malagasy authorities and
will be exported to Robert Baudy in Florida for captive breeding
as soon as the necessary paperwork is completed. Export of tor
toises from Madagascar is controlled by law, and G. yniphora,
being a threatened species, may not be imported into the United
States without a Federal permit. There are very few of these tor
toises even in museums, and the type specimens in the Paris
Museum are of uncertain origin; they were originally supposed to
have come from some small island near the Comores. Some years
ago there were two live specimens at the ORSTOM Center in Tan-
282
anarive, but these have since expired; however ORSTOM is

reported still to have a group of 38 G. radiata.
James Juvik, of the University of Hawaii, recently had the op
portunity of visiting the one tiny area of Madagascar where yni
phora still survives-the region around Cape Sada, near the town
of Soalala, just east of Baly Bay. Juvik eventually succeeded in
locating eight specimens of the ‘Angonoka,’ as the local Sakalav
people call this tortoise, ranging from a four-inch juvenile to two
large adult males about sixteen inches long. All but the two smal
lest specimens were deposited in the Honolulu Zoo in the hope that
they would breed as successfully as the zoo’s Galapagos tortoise
colony had.
The rare Madagascan Geochelone yniphora. Photo by Luis Sosa at the San
Antonio Zoo.
The undeniable extreme rarity of yniphora is not immediately

explicable. The species does not appear to be used for human con
sumption, and although bush fires may destroy some adult tor
toises, and feral pigs eat some of the eggs and young, these factors
m themselves seem inadequate to account for the rarity of the spe-
283
www.Ebook777.com
284
Rhinoclemys rubida perixantha, a terrestrial emydid turtle from Colima, Mex

ico.
Rhinoclemys funerea from Costa Rica.
285
www.Ebook777.com
cies. Juvik has advanced the possibility that the angonoka is a sort
of Irish elk or sabre-toothed tiger among tortoises, a species that
has become so specialized—in this case in the direction of the
greatly elongate gular prong—that the animal is becoming unfit to
survive even in its natural habitat. Juvik observed that the upward
curve of the gular in male specimens appeared to interfere with
the head and neck mobility, and hence, by inference, with feeding
efficiency; moreover the projection appeared too slender and brit
tle to be of use in fighting, and indeed specimens confined together
showed none of the aggressive propensities that similarly equip
ped tortoises such as Chersina angulata display in captivity.
At least one of the angonokas captured by Juvik showed trian
gular dark areas on the carapace scutes, suggesting the pattern of
G. radiata. The occasional specimen of radiata too shows a single
gular scute, so the close relationship between these two species
seems undeniable.
A species possibly ancestral to Geochelone radiata is the ex
tinct G. abrupta, described by Alfred Grandidier, the famous ex
plorer of Madagascar. Remains of at least three individuals have
been found at Amboulitsate; these are sufficient to demonstrate
that the animal was gigantic (at least three feet long), and with
such an elevated shell that the costal scutes were almost vertical.
The species has an amphicoelous (doubly concave-ended) fourth
cervical vertebra, a feature shared with the Galapagos tortoises
but contrasting with the other Indian Ocean giants.
The true giant tortoises of the Indian Ocean islands are classi
fied in two subgenera: the now entirely extinct Cylindraspis, iden
tified by the single gular and absent nuchal, and the living Alda-
brachelys, which normally has paired gulars and a nuchal, and
also a bulbous-fronted, convex-topped skull with an expanded
nasal chamber.
SUBGENUS ALDABRACHELYS
Aldabrachelys originally occurred in Madagascar, the Sey
chelles, Farquhar Island, and Aldabra Island; giant tortoises, pos
sibly of this subgenus, have also been reported from Agalega
Island, the Amirante, Alphonse, and African Islands, and from
Astove Island, but there are no known museum specimens from
these islands in existence. Today the only surviving indigenous
population is on Aldabra Island.
The single Malagasy species, Geochelone (Aldabrachelys)
grandidieri, is known from several nicely preserved carapaces as
well as other, more fragmentary, remains. The size was gigantic
(one carapace in the Paris Museum is about 48 inches long), and
the vault of the shell was up to an inch thick, many times the nor
mal thickness of a giant tortoise shell. The carapace was depres
sed, so much so that this species was originally thought to be a
286
huge freshwater turtle of the family Emydidae. The limbs show

fusion of the carpal elements to an extraordinary degree. The date
of the final extinction of this form is not known; the skeletal
material on hand is true bone rather than fossil, but was mostly
found in dry, sandy caves, under which conditions the bones could
last for centuries without disintegrating. The extinction was
almost certainly not due to human action, but rather to the same
poorly-understood forces which exterminated so much of Mada
gascar’s striking and frequently gigantic Pleistocene fauna.
It is not usually realized that until the end of the eighteenth
century indigenous giant tortoises were found on most of the Sey
chelles Islands. Research into the eighteenth and nineteenth cen
tury writings of such authors as Malavois, Degrandpre, d’Unien-
ville, and Froberville reveals such statements of the tortoises’
abundance as ‘exceedingly numerous’ on Mahe; ‘prodigious num
bers’ on La Digue; ‘a very great number’ on Moyenne; ‘prodigious
numbers’ on Conception; ‘plentiful’ on Recif; and so on down to ‘a
few small ones’ on Marianne. The tortoises have now been eaten
out of existence, and the only museum specimens of even probable
Seychelles origin are the famous large male in the British
Museum, which lived on Mauritius for 152 years before its acciden
tal death in 1918, and possibly a couple of others of similar proven
ance in the Rothschild collection, now housed at Acton. These tor
toises have been called Geochelone sumeirei. They agree with the
Galapagos forms in having paired gulars and no nuchal, but it
must be remembered that a small percentage of Aldabra tortoises
also lack the nuchal. The species is commonly called Marion’s tor
toise, after Chevalier Marion de Fresne, who brought five of them,
apparently from the Seychelles, to Mauritius in 1766 as a goodwill
gift to the Port Louis Garrison.
Farquhar Island is reported once to have supported a large
population of very big tortoises, but only one museum specimen,
assigned to this island on very circumstantial evidence, is known
to exist (presently in the British Museum storehouse at Acton, with
a cast in the California Academy of Sciences). This is a very large
tortoise, 49.50 inches in straight length, similar to G. gigantea but
having a relatively small head and thick, protruding scales on the
head and limbs. The species Geochelone gouffei is founded on this
specimen.
Of all the Indian Ocean giant tortoises, the population of Alda
bra Island, between the coast of Kenya and the northern tip of
Madagascar, is the only one that survives to the present day; all
the others had been wiped out by the hand of man before 1800. The
survival of the Aldabra tortoises may be attributed to several fac
tors, among which are the fact that Aldabra, unlike Mauritius,
Reunion, etc. was a long way off the direct shipping lane from the
287
www.Ebook777.com
Hatchling Rhinoclemys armulata, a terrestrial emydid from Panama and Col
ombia.
Juvenile Rhinoclemys melanosterna , a semi-terrestrial emydid found in

Panama and western Colombia.
288
A pond turtle, Mauremys caspica rivulata, from southern Turkey.
The European pond turtle, Emys orbicularis.
289
www.Ebook777.com
Cape of Good Hope to India and the Orient. Also, the terrain of
Aldabra is so inhospitable that there has never been more than the
most rudimentary settlement on the island, while most of the Mas-
carene Islands have extremely dense human populations.
Aldabra is a coral atoll about eighteen miles long, bordered by
jagged limestone and small beaches and enclosing a large, man
grove-bordered lagoon. The lagoon connects with the ocean in four
places, so that Aldabra is In actual fact composed of four islands-
South Island (the largest); North, or Middle, Island; Polymnie;
and the small Settlement Island, or West Island. Tortoises are
most plentiful on South Island, but a few live on North and West
Islands. The older books list five species of tortoise from Aldabra:
Testudo gigantea, T. elephantina, T. hololissa, T. ponderosa, and
T. daudinii (this was at a time when all tortoises were called “Tes-
tudo,” and the subspecies concept was imprecise and little used;
all must now be considered under the name Geochelone, subgenus
Aldabrachelys). Recent work demonstrates that only one kind of
tortoise is found on Aldabra; it must be called by the oldest avail
able name, G. gigantea. The type specimen of T. elephantina dif
fered from the type of gigantea only in having a single supracaudal
scute; Aldabra tortoises can have a single, asymmetrically divi
ded, or symmetrically divided supracaudal. The single and double
supracaudal conditions are in fact about equally common. T. holo
lissa and T. ponderosa were based on very small numbers of speci
mens differing only in minor details of shape and shell thickness.
T. daudinii was based on a more distinctive specimen with an elon
gate shell and reverted marginal scutes.
Typically, the Aldabra tortoise has a much higher carapace
than its extinct Malagasy relative G. grandidieri. The shell is also
much thinner than that of grandidieri, but nevertheless it is much
thicker than that of most of the Galapagos tortoises. Almost all
specimens have a nuchal scute, paired gulars, and an overall uni
form dark grey or black color. The neck is long and the head rather
small. The area above the eyes is strikingly bulbous and is covered
with two large elongate scales. The snout is somewhat pointed and
the nostrils perforate below the tip; the distance from the nostrils
to the mouth is much less than in a Galapagos tortoise of compara
ble size. The edges of the jaws are finely serrated.
The Aldabra tortoises were apparently quite numerous in the
first half of the nineteenth century, though according to the Rodatz
brothers they kept to the thick scrub. The Rodatz brothers also
mentioned in their report that they had found two brick-walled
enclosures on the island, housing 200 and 300 tortoises respective
ly ; these presumably had been get together by a shore party for
stocking ships. According to Kirsten, who quotes a Hamburg mer
chant, 1200 tortoises were removed from Aldabra in 1847. By the
290
end of the nineteenth century, the tortoises were apparently rare;

the German traveller Voeltzkow made two trips to Aldabra in the
1890’s, finding only two tortoises on the first trip and seven on the
second. Voeltzkow’s account of his second voyage in 1895, quoted
by Gadow, includes the following passage:
“The chief hindrance in the search for tortoises is the
inpenetrability of the island. The soil consists entirely of
sharp water-worn corals, with their points uppermost, while
the whole is covered ivith such thick masses of low scrub,
that a way has to be cut with an axe, so that an extended
search over a large area is out of the question. To land on the
outside is dangerous, on account of the heavy surf; while
landing from the inside of the atoll is much hindered by the
dense thickets of mangrove trees. As drinking water, and
that very bad, is only found in one place, rainwater has to be
collected from the natural hollows, and carried along in
tanks. Thousands of mosquitoes prevent one remaining over
night in those places which the tortoises frequent. Then at
last, when one has discovered, by a stroke of luck, one of
these creatures, the real hard work begins, namely the con
veyance of the beast. ”
Somewhat prior to Voeltzkow’s visits, in 1874, anxiety had
been felt for the future of the Aldabra tortoises when it was an
nounced that a permanent party of woodcutters was to be placed
on the island. A formal petition, signed by such eminent natural
ists of the day as Charles Darwin, Joseph Hooker, H.B.E. Frene,
Richard Owen, Alfred Newton, John Kirk, and Albert Gunther,
was submitted to the Governor of Mauritius, to the effect that a
number of tortoises should be transferred to Mauritius and the
Seychelles, where they could breed in captivity and save the spe
cies should those tortoises remaining on Aldabra all be killed. The
Governor followed this suggestion, and some of the tortoises were
transferred to the Seychelles, where a breeding colony became es
tablished.
Judging by the apparent decline of the Aldabra tortoise popu
lations during the nineteenth century, one would imagine that to
day they would be virtually extinct. It is surprising, therefore, to
read recent accounts of Aldabra, such as those of Ommaney, Gay-
mer, Grubb, and Frazier, which record the astonishing abundance
of tortoises on the island today; around water holes it is possible to
include fifty tortoises in a single photograph, and total population
estimates run from 33,000 to over 100,000-the latter estimate com
municated to me by Jack Frazier, who spent ten months on the
island. The island, in fact, gives every appearance of being over
run with tortoises, and the species has been removed from the Red
Data Book of threatened spe cies. The reasons for this apparent
291
www.Ebook777.com
Mauremys mutica, a rather small emydid turtle from Taiwan, Hainan,
southern China, Japan, Vietnam, and the Ryukyu Islands.
292
a colorful but relatively rare Asian emydid. Above, heads of

Sacalia bealei,
male (left) and female (right); below, plastron.
293
www.Ebook777.com
Large number of Geochelone gigantea at a pool west of Cinq Cases, Aldabra.
Photo by J. Frazier.
population explosion of tortoises during the present century are

unknown; it is also not certain if the tortoises are within the perm
anent carrying capacity of the habitat, Aldabra being a small, and
island, while tortoises have very large appetites. On the other
hand, old reports suggest that the Mascarene Islands carried a
similar density of tortoises until the early eighteenth century.
There is some suggestion that the Aldabra tortoises are not repro
ducing to an extent even remotely comparable to their numbers,
Gaymer found very few young tortoises in 1964 and 1965, and more
recently Frazier, during ten months on the island, saw tortoises
copulating on very few occasions, and actual nesting only three or
four times; moreover, he found very few completed nests, even
though the nest site, which the female drags herself over for days
after laying, remains obvious for some time. It is possible that toe
tortoises are not receiving an optimal level of nutrition and that by
some built-in mechanism their reproductive instincts are being in
hibited' it is also possible that copulation is inhbited simply by the
over crowding on the island. Keepers of captive giant tortoises ob
serve much more reproductive activity when the sexes are kept
apart for some time and then are re-introduced to each other. Pos
sibly the constant availability of mates in the wild has a similar
inhibitory effect.
294
As with the other large tortoise species, male Aldabra tor

toises are substantially bigger than the females; the average adult
female is about thirty inches long, while all individuals more than
forty inches long are males. Some tortoises of only medium size
appear to be very old, judging by their smooth, worn-down shells,
while on the other hand the largest individuals sometimes appear
to be relatively young. The young have the potential to grow very
rapidly, which is a necessary adaptation since the tortoises less
than eight inches long are liable to be preyed upon by several large
bird species, as well as by land crabs. It is estimated that the tor
toises in the wild take about 20-25 years to reach maturity; males
go on growing about five years longer than females. The growth is
very uniform in the early stages, but later on individual differen
ces appear; some tortoises remain rather smoothly-domed, while
the scutes of others are distorted into conical bosses; the latter
condition may possibly be a result of especially rapid growth.
When large size is reached, growth becomes very slow; the actual
direction of post-maturity growth is very variable, and the dif
ferent appearance of very large specimens has led to some taxo
nomic confusion. Many old males develop strikingly swollen
bridges.
The daily life of tortoises on Aldabra involves long periods of
lying in mud wallows and freshwater ponds; the primary purpose
of this is presumably thermoregulatory, though other factors may
also be involved. The tortoises start feeding early in the morning
and continue until the sun becomes too hot; by 9 a.m. they are usu
ally obliged to seek shade. The tortoises are mainly vegetarian,
feeding largely on dry grasses and dead leaves, as well as Ficus,
Scaevola, and Sporobolus. However, they will eat meat when it is
available, and Gaymer reports local stories of cannibalism of dead
tortoises on Aldabra, and even of a captive tortoise in the Seychel
les which learned to kill piglets and chickens by letting them move
beneath its raised plastron and crushing them by suddenly with
drawing its legs; the animals were then ground to pieces and
eaten.
The tortoises are very shy in the wild and, while they can be
observed from very close range if one keeps behind the rise of the
shell, as soon as the tortoise spots the observer, the head and limbs
are pulled in with a loud, prolonged hiss, and the animal sinks to
the ground as if struck dead.
As was mentioned earlier, Aldabra tortoises nowadays seem
to show amazingly little breeding activity, and quantitative data
on clutch size and other aspects of breeding biology in the wild are
not available. Moreover, zoo specimens, although they mate regu
larly, almost never seem to lay eggs, and the Aldabra tortoise has
never been bred outside the Indian Ocean Islands. However, semi-
295
www.Ebook777.com
Annamemys annamensis, a rarely seen emydid from Vietnam.
Batagur baska, a giant emydid river turtle from the Malay Peninsula.
297
www.Ebook777.com
captive tortoises on various islands in the Seychelles have been
breeding regularly for many decades, and a number of interesting
observations have been made on the behavior of the herd of Alda-
bra tortoises in the pare aux tortues in the Seychelles. The breed
ing season extends from January to April; the females carry the
eggs for about ten weeks, after which period they are buried in the
ground in the normal tortoise fashion. Between 9 and 25 eggs are
laid at a time (considerably more than were present in the few
nests found on Aldabra itself by Jack Frazier); they are white,
spherical, and the size of tennis balls. They take 120 to 130 days to
hatch. As in the Galapagos, when the ground is very hard the
young may have difficulty emerging. The fertility rate is about
50%. One female may nest twice in a season. After four years, the
baby tortoises reach a length of 18 to 22 inches; on Aldabra itself,
on the other hand, where food and water are scarce for much of the
year, four year old tortoises are only eight or nine inches long.
Both in the wild and in captivity, however, full growth appears to
be reached after about 25 years. In the period 1904-1909, the captive
herd produced 168 young. Forty-nine of the tortoises, including 18
mature females, were released on Long Island, in the Seychelles,
in 1910. It was formerly a local custom in the Seychelles to mark
off a young tortoise when a child was bom and to eat that tortoise
on the child’s wedding day.
From time to time, rumors are propagated of some of the in
digenous Seychelles tortoises having survived to the present day.
However, it appears that these accounts are based either on cap
tive-raised Aldabra tortoises with the minor shell deformities that
often develop under captive conditions, or on Aldabra tortoises
which, by way of individual variation, lack the nuchal scute (Gay-
mer found that three Aldabra tortoises out of 223 examined lacked
this scute).
Several individual Aldabra tortoises have achieved a degree of
fame, usually on account of their longevity. Perhaps the best
known is Jonathan, resident in the grounds of Government House
on St. Helena Island in the South Atlantic, and by popular repute a
contemporary of Napoleon during his exile and last years on the
island. Unfortunately for this attractive story, it is now known that
Jonathan was imported in 1882 and thus missed Napoleon by 61
years. However, a female tortoise, almost certainly of Aldabran
origin, was brought to St. Helena via Mauritius in or around 1776,
and a second specimen, supposedly a male but in fact a female,
was presented to the Governor of the colony by Captain James
Pritchard around 1826. The first of these tortoises died in 1877, and
the second in 1918, following a fall down a steep slope. A third tor
toise, the one still living on the island, arrived in 1882, as a replace
ment for the tortoise which had died five years before. This tor-
298
toise was christened Jonathan by Governor Sir Spencer Davis dur

ing his four-year term 1932-1936. By 1969, Jonathan was showing
definite signs of extreme age, so two more Aldabra tortoises, each
about 10.5 inches long, were sent to St. Helena in that year by Sir
Bruce Greatbatch, Governor of the Seychelles.
Apparently the largest Aldabra tortoise of which there is a
definite record, and perhaps the largest modem tortoise known,
was one which lived on Egmont Island, in the Chagos Archipelago,
for many years. The tortoise eventually passed into the hands of
Lord Rothschild, who deposited it in the London Zoo, where it died
at the beginning of the present century. The specimen is now
mounted in the British Museum; it has a straight-line carapace
length of 55 inches and weighed 560 pounds even when in poor con
dition after transportation to Britain.
Another famous tortoise was brought to Colombo, Ceylon, by
the Dutch, and was found there by the British in 1796. It died in
1894, a week after being moved to new quarters in Victoria Park.
Another very large Aldabra tortoise, with a 53.5 inch carapace,*
was still alive at Hirimbura, Garstin Hill, Galle, Ceylon, in 1910;
evidence that is now lost suggested that this tortoise had been
brought to Ceylon in 1797.
Specimen of the Aldabra tortoise, Geochelone gigantea, showing the

distorted carapace scutes often found in long-term captives. Specimen in the
London Zoo.
www.Ebook777.com
Kachuga tentoria circumdata, a newly described subspecies of the Indian
roofed turtle from the Calcutta region.
www.Ebook777.com
One male and three female Aldabra tortoises were brought to
Ceylon by Sir Reginald Stubbs in 1936. This group produced
several clutches of eggs, which were reportedly 49-50 mm in diam
eter and thus smaller than the ‘tennis ball’ sized eggs laid by Alda
bra tortoises in the Seychelles. Nine hatchlings were produced, one
of which lacked a nuchal scute. The hatchling carapace lengths
ranged from 62 to 69 mm, and the weights from 40 to 48 gm (when
two days old). The smallest tortoise reached a length of 764 mm
and a weight of 220 lbs. in 18 years and 7 months.
SUBGENUS CYLINDRASPIS
The tortoises of the Mascarene Islands are now entirely ex
tinct, and have been so since about 1800. About a dozen species
have been described, all originally under the genus Testudo, but
now considered to belong to the subgenus Cylindraspis of the genus
Geochelone. The species Testudo vosmaeri, T. peltastes, T. com-
mersoni, and T. rodericensis, were described from Rodriguez;
Testudo indica from Reunion; Testudo inepta, T. leptocnemis, T.
triserrata, and T. gadowi from Mauritius; and T. microtympan
um, T. sauzieri, T. grayi, and T. africana, of doubtful origin and
highly questionable distinctness. It is in fact very probable that
each island harbored only a single, very variable form. For exam
ple, T. rodericensis is a straight synonym of T. vosmaeri; T. pel-
tastes is based on some rather small shells from Rodriguez, but
they are probably either juveniles or females of T. vosmaeri. No
type specimen of T. commersoni exists, the species being based
entirely on a drawing.
The first giant tortoise to be brought to the attention of zoolo
gists was Geochelone (Cylindraspis) indica from Reunion, of
which a shell three feet long, now lacking the plastron, arrived in
Paris in 1676, reportedly from the coast of Coromandel. Perrault
named it “Tortue des Indes” (this was before the binomial system
of nomenclature was instigated). Linnaeus completely overlooked
this specimen, which had to wait until 1783 for a scientific name,
when Schneider called it T. indica. Later, Dumeril and Bibron re
named this specimen Testudo perraulti. The form is characterized
by the elevated front of the carapace, the margin of which is some
what reverted, very much in the fashion of some of the saddleback
tortoises of the Galapagos. The hind margin of the carapace is
strongly serrated. As with all the subgenus, the nuchal is absent
and the gular scute is small and single. The species was at one
time very abundant, but it was the first of the Mascarene tortoises
to become extinct. By the mid-eighteenth century not one was left,
the result of the usual factors of over-exploitation by visiting ships,
cultivation of the land with consequent habitat destruction, and in
troduction of feral mammals which either competed with the tor
toises for food or destroyed the eggs and young.
302
The tortoises of Rodriguez, too, were extraordinarily abun

dant in the seventeenth century. Francois Leguat, who went to the
island in 1691, published an account of his findings in several lan
guages in 1708; the section pertaining to the tortoises is as follows:
“There are no four-footed animals at Rodriguez, hut
rats, lizards, and the land turtle. Of the latter there are three
different kinds; and some of them are so large as to weigh an
hundred pounds. Their flesh is wholesome, and not unlike
our mutton, but more delicate. The fat, which is white, does
not congeal, and is never known to cause an indigestion; we
consider it as superior to the finest butter in Europe. The
liver is a most delicious food, and is very large in proportion
to the animal, as it is equal to one-third of the whole weight
of the flesh. The bones are solid, and consequently without
marrow.
“These turtles lays their eggs in the sand, with which
they cover them; when they are left to the slow operation of
being hatched by the sun. They are perfectly round, like bil
liard balls, and are not much larger than those of our com
mon fowls. The shell is soft, and the interior part is excellent
food.
“There is such an abundance of these land turtles, that
they are sometimes seen in flocks of two or three thousand.
They collect together towards the evening, in the coolest
places they can find, and in such close array, that the spot
which they have chosen seems to be paved with their shells.
It has been observed that some of them constantly take their
post at a small distance from the main body, as if to perform
the duty of sentinels; but the utility of this arrangement is
not readily comprehended, as the turtle possesses no means
of active defense, or the least capability to save itself from
danger by flight. ”
It is interesting that Leguat mentions the existence of three

kinds of tortoise on Rodriguez, but since he also described the eggs
as being soft-shelled and being laid in the sand, it is likely that his
account is a composite description of the land tortoise and two spe
cies of sea turtle (probably the green and the hawksbill). To com
plicate matters further, the very existence of Francois Leguat has
been called in question by G. Atkinson in 1922, who suggested that
Leguat’s supposed work was in fact a novel written by F.M. Mis-
son. However, this theory is fairly satisfactorily laid to rest by G.
Evelyn Hutchinson in his book The Enchanged Voyage.
Another account of the Rodriguez tortoises, written by the
Mauritius magistrate M. Rouillard between 1730 and 1760, includes
the section:
303
www.Ebook777.com
304
Plastral view of Heosemys grandis.
OPPOSITE: Two views of the seldom seen semi-terrestrial Geoemyda

from Southeast Asia. Photos by Dr. Robert Mertens.
spengleri
305
www.Ebook777.com
“The land tortoise is very abundant. It is not very fat, I
owing to the great number of them and the dearth of grass;
it eats leaves and the fruit of the trees, which the wind
causes to drop to the ground. There are tortoises of three
species, and the largest which I have seen are from three
feet to three feet eight inches in length of shell. They are not
so common in the heights as in the ravines, on account of the
dearth of water in the dry seasons. ”
The mention of three species may be an uncritical quote from
Leguat, since the simultaneous existence of three separate forms
of giant tortoise on such a small oceanic island is zoogeographical-
ly and ecologically unlikely. The great abundance of the tortoises,
and their lean condition, appears to have been the primordial con
dition on all of the tortoise islands of the Indian Ocean, and such a
situation apparently exists on Aldabra today.
By the mid-eighteenth century the tortoises of the neighboring
island of Mauritius had become scarce, and large numbers were
imported from Rodriguez; in an 18-month period in 1759-1760,
30,000 tortoises were imported from Rodriguez, and many more
were removed unofficially. The majority of these did not survive
the trip to Mauritius-a typical log book entry is as follows;
“1759, 16 Decbr., the Penelope arrives from Rodriguez
with 1035 land tortoises and 47 turtles. The cargo was of 5000
of the former and 50 of the latter; but a passage of eight days
reduced the number to the few which she brings. ”
The massive mortality of the tortoises compared to the turtles
is probably attributable to their exceedingly thin shells, which
cracked easily when the tortoises were piled on top of each other in
the holds.
As early as 1726 or 1727, a plea for a more sane approach to the
exploitation of Rodriguez tortoises was raised by one M. Lenoir,
who wrote in a latter to the council of the French East India Com
pany:
“It cannot be permitted that ships going to the Indes and
returning there, should go without hindrance to despoil the
neighboring islets of the land-tortoises; and it is necessary to
prohibit captains from sending their boats to take them un
less the commandant of the island gives permission, and
gives the number which they may take away. ”
Sadly, this voice of reason was ignored completely. By 1761 the
tortoise populations had been reduced substantially, and by 1770
they were so rare that a trading station established in the interior
of Rodriguez as a collecting center for tortoises was abandoned.
Their extermination was completed in the next twenty years, in
part by the imported cattle and pigs which destroyed the eggs and
306
trampled or ate the young of the last few tortoises. In 1864 only a
few crumbling shell fragments could be found.
The Rodriguez tortoise was officially introduced to science by
Schoepff, who called it Testudo indica vosmaeri. Schoepff’s de
scription was based on a shell, reputedly from the Cape of Good
Hope, one of the very few in existence collected during the lifetime
of the species and thus with the scutes intact. This shell is now in
the Leiden Museum; it is 32 inches long and 18.5. inches wide. Ori
ginally the carapace was described as black and the plastron as
yellowish. This coloration is unknown among living giant tortoises,
which are black both above and below. The Leiden specimen too is
now completely black, but it appears to have been painted with
some sort of black paint or preservative which has obscured the
original coloration.
The Rodriguez tortoise on occasion reached a very large size;
some of the bones in the British Museum, according to Gunther,
correspond to individuals about 4.5 feet in length, though other ap
parently mature specimens were much smaller. The shell is ex
ceedingly thin, and the animal had the long neck and limbs and
raised front of some of the Galapagos tortoises. The plastron was
very small, strongly narrowed in front and shortened at the rear.
The shape is somewhat similar to that of the Reunion tortoise, but
the carapace margin is less wavy, the three tubercles on the front
carapace scutes are lacking, and the carapace margin is nowhere
reverted.
Geochelone vosmaeri is also known from two shells with
scutes in the Paris Museum and a complete stuffed (though rather
distorted) male, located some years ago in the natural history col
lection of the Library of Saint Genevieve, and now also in Paris (a
plaster cast is in the British Museum). Numerous bones, mostly
fragmentary, are distributed among several European museums.
Bones corresponding to several hundred individuals were collected
by an astronommical expedition (the so-called “Transit of Venus
Expedition”) in 1876 and were deposited in the British Museum.
A similar story of over-exploitation resulting in complete ex
tinction could be told for the tortoises of Mauritius. Apparently
they were not as large as those of Rodriguez, few specimens ex
ceeding a carapace length of thirty inches, but they made just as
Outline of shell of Geochelone

vosmaeri.
307
www.Ebook777.com
Two views of the spiny turtle of Southeast Asia, Heosemys spinosa. Com
pare with H. grandis on a previous page.
The Ceylon terrapin, Melanochelys trijuga thermalis; younger specimens

have bright yellow or orange spots on the head.
Melanochelys trijuga trijuga, an Indian emydid turtle.
309
www.Ebook777.com
good eating. No less than seven scientific names have been applied
to the tortoises of Mauritius: Testudo gadowi, T. inepta, T. lepto-
cnemis, T. triserrata, T. boutonii, T. africana, and T. grayi. In
addition, two other names, Testudo sauzieri and T. micros
tympanum, may be based upon material from Mauritius.
Testudo gadowi (now, of course, properly known as Geoche-
lone gadowi) was a much earlier form than the others, which be
came extinct well before the discovery of the island by man and is
easily distinguished by its elongated and forked gular area. Such a
modification is found repeatedly in the Testudinidae (e.g. in Goph-
erus berlandieri or in the extinct Geochelone (Megalochelys) siva-
lensis), but this is the only known case of its occurrence in a Mas-
carene tortoise, all others having only a single, very small, gular
scute.
Of the remaining names, we can immediately synonymize
africana and grayi since they are based upon the same specimen,
a sixteen-inch carapace, still with scutes intact, in the Paris Mu
seum. The history of the specimen is not known, but Gunther con
cluded from the undulated vertebral profile that the animal was
originally from Mauritius, and if this is so it is the only known spe
cimen of a Mauritius tortoise with the scutes still in place. It was
originally named Testudo tabulata var. Africana (i.e. as a race of
a South American tortoise!) by Schweigger in 1812, but this name
seeming inappropriate, it was named Testudo grayi by Dumeril
and Bibron in 1835.
Testudo boutonii is based upon a single scapula, and can be
dismissed from further discussion, as can Testudo sauzieri and T.
microtympanum, since neither sufficient data nor specimens are
available to allocate these forms definitely. We are left then with
T. inepta, T. leptocnemis, and T. triserrata. T. triserrata differs
from T. inepta in having an extra cutting ridge on the lower jaw;
but since the remnants of this tortoise were found indiscriminately
mixed with those of inepta in a bog known as “Mare aux Songes,”
it seems very probable that the Mauritius tortoise was polymor
phic in respect to the number of jaw ridges. The bones of T. leptoc
nemis, characterized by their slender, gracile form, were found in
a different part of the island, in the District of Flacq. Possibly dif
ferent environmental conditions in what are now the Mare Aux
Songes and the District of Flacq caused differences in the massive
ness of the bones of the tortoises.
The few intact carapaces of the Mauritius tortoise in the British
Museum and elsewhere show that this form was smaller than the
Rodriguez species, with a thicker shell. The carapace lacked the
elevated front margin of the Rodriguez tortoise and had a some
what wavy vertebral profile, with a particularly deep indentation
at the junction of the fourth and fifth vertebral scutes.
310
SUBGENUS MANOURIA
The two living Asiatic tortoise species included in the sub
genus Manouria are considered to be the most primitive living tor
toises, very close to the ancestral Eocene stem which later gave
rise to such genera as Gopherus and such subgenera as Indotes-
tudo and Chelonoidis. In fact, Auffenberg (1971: Copeia, p. 106)
has recently made the suggestion that the primitive tortoises of the
subgenus Hadrianus are not separable subgenerically from
modern Manouria. The primitive features of the subgenus, which
betray its closeness to the ancestral emydines, include the broad,
flattened shell, the wide, posteriorly expanded nuchal scute, the
paired supracaudal scutes, the lack of any excavation inside the
gular lip of the plastron, and the preference for mesic, rather than
xeric, habitat.
The Burmese brown tortoise, Geochelone (Manouria) emys, is
found in Assam, Burma, Thailand, the Malay Peninsula, Sumatra,
and Borneo. It is the largest of the Asiatic tortoises, reaching a
carapace length of at least eighteen inches. The broad, low shell is
of a rich brown to black color; the nuchal scute is well developed,
Plastral view of the Burmese brown tortoise, Geochelone emys. Notice the
widely separated pectoral scutes.
311
www.Ebook777.com
Cyclemys mouhoti, a largely terrestrial emydid from Vietnam, is often as
signed to the genus Pyxidea.
The Asiatic leaf turtle, Cyclemys dentata.

www.Ebook777.com
the supracaudals are paired, and the vertebral region is flattened.
There is often a shallow concavity in each of the costal scutes. The
plastron is less strongly pigmented than the carapace, and the pec
toral scutes in most specimens are completely and often widely
separated. The gular scutes are extended forward. The head is of
moderate size, light to dark brown in color. There is a large frontal
scale and two pairs of prefrontals. The faces of the forelimbs are
covered with thick, heavy, pointed, overlapping scales. There is a
patch of tubercular scales on each thigh so large that this species
is often called the six-footed tortoise.
This highland species is becoming scarce in most parts of its
range, since it is eaten almost whenever found. According to Mal
colm Smith, the eggs measure 43 by 50 mm, but otherwise no data
are available on any aspect of its reproductive habits.
The other species of Manouria is Geochelone impressa, the im
pressed tortoise. This is a much smaller species than G. emys, not
exceeding a carapace length of about 10 or 11 inches. Its range is
very similar to that of G. emys, except that it does not extend as
far north or south, being absent from Assam, Sumatra, and Bor
neo. This too is a hill species, always being found at considerable
altitudes. The carapace is sometimes almost uniform brown, but
in other specimens the rich brown ground color is marked with
striking black edgings and radiations on each scute. The vertebral
region is very flattened, and the costal scutes are distinctly con
cave. Both anterior and posterior carapace margins—and often
the sides—are strongly serrated and slightly reverted. The head is
of moderate size and somewhat pointed, and the nasal region is
often pink. It is not known if this pink coloration is seasonal. The
limbs are covered with thick, heavy scales. In contrast to G. emys,
the pectoral scutes always have a strong median line of contact.
The gula'r scutes are produced forward and separated anteriorly;
there is a deep notch in the rear of the plastron. Juveniles are of
similar shape to the adults, but differ strikingly in having fine
black specks all over the carapace and head.
The impressed tortoise is apparently a rare species, almost
never seen in captivity, and its habits are almost entirely un
known.
SUBGENUS INDOTESTUDO
The third Asiatic subgenus is Indotestudo, a rather general
ized group normally considered to consist of three species which
are restricted to mesic tropical forests of southeast Asia. The sub
genus is characterized by having the humeropectoral sulcus cross
the rear part of the entoplastron; otherwise the characters are
mostly those typical of the genus as a whole: the nuchal scute may
or may not be present; the supracaudal is undivided; the gular
scutes are paired but not elongated; the pectorals are not nar-
314
The Southeast Asian Geochelone elongata is fairly distinctive in having a

whitish head that becomes partially pink during breeding.
rowed; and the neural bones show an alternating arrangement of

octagonal and square bones. As with many forest-living tortoises,
the carapace is relatively narrow.
Geochelone elongata is found from north-eastern India
through Burma, Thailand, Laos, and Cambodia, east to North Viet
Nam, and south as far as Penang, Malaysia. It has a rather nar
row, elongate carapace, basically unpigmented, but with smudgy
black spots on the costals and centrals and on most of the margi
nals. The plastron similarly is basically unpigmented, but has a
large black blotch on each of the abdominal scutes, and often also
on the humerals, pectorals, and femorals. A small, narrow nuchal
scute is almost always present. In adult specimens the interpec
toral seam is slightly longer than the interhumeral seam. The head
scalation consists of a well-defined pair of prefrontals, a single
frontal, a small occipital, and various dorsolateral and lateral
scales of variable size. The jaw surfaces are feebly denticulate,
and the upper jaw is tricuspid. The head is unpigmented, appear
ing pale greenish-yellow or almost white. In the breeding season
the skin around the nose and eyes becomes pink, hence the Bur
mese name for the species (“laik nakhonga”—red-nosed tortoise).
315
www.Ebook777.com
Juvenile Notochelys platynota, a semi-terrestrial Asiatic emydid.
Cuora flavomarginata, a box turtle abundant on Taiwan, where it is often

called the snake-eating turtle.
317
www.Ebook777.com
It is possible that the pink color results from an increased vascu
larization of those sensory systems associated with sex and
species recognition—i.e., the olfactory and visual systems. The
limbs are greyish, with large flat scales, especially along the outer
edge of the forelimbs and on the heels.
The habits of this species have not been studied. It is usually
found in hilly or mountainous areas, possibly in part because those
individuals in the more low-lying areas are hunted for food. It
appears to be much more abundant in the southern part of its
range than in the northern part. According to K.G. Gairdner (quo
ted in Malcolm Smith: “Fauna of British India,” Vol. 1,1931), this
species has a remarkable tolerance for extreme heat, being found
on open hillsides in Thailand when the ground was so hot that the
hand could hardly bear to touch it. Nevertheless its typical habitat
is mesic rather than xeric. By preference this species pursues a
frugivorous diet.
The Travancore tortoise, Geochelone travancorica, is restric
ted to hill country in the southern tip of India, in the State of Ker
ala ; it is most plentiful on the western slopes of the western Ghats,
but has also been recorded from the eastern slopes. It is very simi
lar to the elongate tortoise, from which it has probably been sepa
rated since the late Pleistocene, but differs in that it always lacks
the nuchal scute, has a somewhat less elongate carapace of a
darker, browner overall color (though still with large black
blotches on each scute), and the interpectoral seam is shorter than
the interhumeral, instead of the reverse.
The habits of the Travancore tortoise have received little at
tention, but the courtship patterns have been studied by Auffen-
berg. This species, like elongata, assumes a reddish color around
the nostrils and eyes during the breeding season (November to
January). The sex of the intended mate is confirmed by smelling
the cloacal region by means of a characteristic movement involv
ing extension of the head, followed by rapid movement of the head
through a short vertical arc; the movement terminates in a small
and rapid curlicue. Following sex recognition, the female is “im
mobilized” by repeated ramming by the male. This is a common
practice among courting tortoises, but it is carried out for particu
larly long periods by this species and may be important not just for
immobilization of the female but also for bringing the hormonal
systems of one or both tortoises to a state of readiness. Actual
mounting follows the variable period of shell ramming; mainten
ance of the mounted posture is probably assisted by insertion of
the claw-like tailtip into a shallow depression at the level of the •
femoro-anal seam of the female.
Copulation with the sympatric Geochelone elegans is probably
avoided not by any complex species recognition ritual, but simply
318
by the relatively flat-plastroned male of travancorica being unable

to maintain its position on the very highly vaulted shell of the
female elegans. Also, in such an attempted mixed copulation, the
tail of the male travancorica would be too far forward to make con
tact with the cloaca of the elegans, and elegans has no femoro-anal
depression to receive the homy tailtip of the male travancorica.
The male Travancore tortoise is relatively silent when mating, but
may produce faint grunts. The head is stretched out and downward
during copulation, and the mouth is held wide open.
The third species of Indotestudo that has been described is
Geochelone forsteni, restricted to the islands of Celebes
(Sulawesi) and nearby Halmahera in Indonesia. Boulenger de
scribed this species as very similar to elongata, only differing in
the lack of a nuchal scute and the shorter interpectoral seam.
Since these are the precise characters in which travancorica dif
fers from elongata, the possibility cannot be overruled that for
steni is based on introduced specimens of travancorica. It is hard
otherwise to rationalize the absence of the subgenus in Borneo.
Wermuth and Mertens (1961: Schildkroten, Krokodile, Brucken-
echsen, p. 188) separate the two species on the basis of the relative
ly broader gular scutes of travancorica and the presence only in
forsteni of a large dark spot on each abdominal scute. However,
the former of these supposed differences is probably the result of
comparing juvenile forsteni with adult travancorica, and the latter
simply does not hold true—most travancorica have black areas on
the abdominal scutes.
Juvenile tortoise refer

red to Geochelone
forsteni.
&§,:-4j
www.Ebook777.com
320
The black pond turtle, Geocleimys hamiltoni.
Chinemys kwangtungensis, an emydid turtle from mainland China.
Reeves’ turtle, Chinemys reevesi.
www.Ebook777.com
SUBGENUS CHELONOIDIS
The genus Geochelone is no longer represented in North Amer
ica, and there are only three species of the genus in South Amer
ica, together with a fourth polytypic species in the Galapagos Is
lands. However, these three species are abundant and very wide
spread, between them covering virtually the entire continent ex
cept for the high altitudes and Patagonia. They constitute a sub
genus, Chelonoidis, which is very closely related to the Asiatic In-
dotestudo and is characterized by the absence of the nuchal scute,
the large entoplastron, the undivided supracaudal scute, the con
cealed postotic fenestra, the relatively wide pectorals, and the vir
tual or complete absence of radiating patterns on the carapace
scutes. Four species of the subgenus Chelonoidis are recognized:
Geochelone denticulata, G. carbonaria, G. chilensis, and G. ele-
phantopus. The sum total of characters suggests that the first two
form a natural group and the last two (despite their extreme dis
parity in adult size) another. The “carbonaria“ group is charac
terized by the thick carapace of a high narrow shape with parallel
or concave sides; the dentate carapace margins of the young; the
narrow posterior plastral lobe; the subtriangular inguinal scute;
the rounded skull roof profile; the small supraoccipital process;
the relatively slight expansion of the ends of the humerus; and the
twisted radius, whose ends are not greatly expanded. The “chilen
sis” group is characterized by the thin, even fenestrated, cara
pace, whose sides are parallel to convex (except in the Galapagos
saddlebacks); the lack of dentations in the carapace margin; the
wide posterior plastral lobe; the anterior narrowing of the inguinal
scute; the flattened or even concave skull roof, with a well-devel
oped supraoccipital crest which rises above the level of the top of
the skull; the great expansions of the ends of the humerus; and the
flattened, terminally expanded radius.
Geochelone denticulata, the yellow-foot tortoise, and G. carbo
naria, the red-foot tortoise, are found sympatrically over much of
their range, which encompasses almost the entire tropical low
lands of South America. The range of denticulata, however, does
not extend west of the Andes and does not enter Paraguay or Rio
de Janeiro State, Brazil, but does extend into north and east Boli
via. The range of carbonaria extends west of the Andes into the
Choco of Colombia and into western Panama, and to the south the
species reaches Rio de Janeiro and Paraguay; but it is not found in
the upper reaches of the Amazon drainage in Amazonas, Peru, and
southeast Colorfibia, or in Bolivia. I believe both species occur
naturally on Trinidad, and carbonaria has reached a number of
other West Indian Islands, presumably by introduction; popula
tions are known on the Grenadines, Barbuda, Old Providence
Island, Long Island (north of Antigua), St. Thomas, Water Island,
322
St. John, Tortola, Peter Island, St. Barthelemy, Montserrat, and

Grenada. There is an introduced colony in the crater of Mount
Misery, St. Kitts.
The yellow-foot tortoise, Geochelone denticulata, is one of the
largest mainland tortoises. Although females are sexually mature
when only about a foot long, the maximum size is much greater
than this. The largest specimen from the Guianas I have seen was
a 24” female; a 22” individual from the Upper Corentijn River
weighed 56 pounds when alive. The species appears to reach a con
sistently large size in Amazonian Peru; two specimens from this
area in the American Museum of Natural History have carapace
lengths of 26.5” and 25.3” respectively. Dick Goergen, of East Am
herst, N.Y., has a 27.75”, 116 pound captive specimen which is still
growing. Moreover, I recently saw a specimen with a 29” carapace
in the Tarpon Zoo, Florida, which had been imported from the
vicinity of Leticia, Colombia, on the Peruvian border. Males are
smaller than females (I once saw a very old 20” male from Guy
ana, but males rarely exceed even 16”) and have a rather different
shell shape; the females have relatively high, though narrow,
shells without a plastral concavity—though the hind lobe of the
plastron is bent somewhat downward. Males have elongate, low
shells with a distinct expansion at the level of the hind feet, and the
plastral concavity may be very deep. The most obvious (though
not necessarily the correct) interpretation of the narrow shell of
adult denticulata is that it presents less of an obstacle when walk
ing between narrowly spaced saplings. In mature animals of both
sexes the shell is very thick and heavy. The overall coloration of
the carapace is dark brown, with a broad, ill-defined yellow-
orange area on each of the scutes. The plastron too is dark brown,
becoming lighter toward the juvenile part of each scute. The ex
tremities are basically brown, with bright orange scales on the
limbs and orange markings on the sides and top of the head.
The yellow-foot tortoise is known as “motelo” in Peru, as
"morrocoy” in Colombia and Trinidad, as “jaboti” in Brazil, and
as “sigrilpatoe” in Surinam. Its preferred habitat is dense rain
forest. Its diet is vegetarian, with a propensity toward fruit; in the
Guianas it tends to frequent the vicinity of wild plum trees. It is
also reported to feed upon leaves and mushrooms. Eggs are repor
tedly deposited in December in the lower Caqueta River, Colom
bia, and in August and September in the Upper Caqueta and Putu
mayo. According to Medem, the eggs may number from 15 to 20
per clutch and are either buried in shallow nests or simply deposi
ted among the dead leaves on the forest floor. I believe the normal
clutch size in the Guianas is much less than this. The eggs are al
most spherical, with a thick, hard shell and are very variable in
diameter.
323
www.Ebook777.com
Adult male spurred tortoise, Geochelone sulcata, the largest mainland tor
toise species.
Baby radiated tortoise, Geochelone radiata, hatched in captivity at the

Gladys Porter Zoo.
Geochelone yniphora, a very rare tortoise from northwestern Madagascar.

Photo by James Juvik.
Comparison of Geochelone gigantea, the only living Indian Ocean giant tor
toise (in background), with the Galapagos giant tortoise. Photo by Dr. Herbert
R. Axelrod.
www.Ebook777.com
Throughout its range the yellow-foot tortoise is esteemed for
its food value, and around human habitations and settlements the
population densities inevitably diminish rapidly. The animals are
typically kept alive for transportation to the markets; in the Guia-
nas, and probably elsewhere, the people take advantage of the
strongly upcurved gular region to wedge a stick horizontally
across the anterior shell opening to keep the animal from extend
ing the head and front legs and so escaping. Frequently a similar
stick is passed across the hind shell opening, and the two sticks
tied together to make a convenient handle. In some parts of Vene
zuela the local people have the destructive habit of setting fire to
whole hillsides so that they may walk through afterward and pick
up the exposed (and presumably often roasted) tortoises. How
ever, the species appears to be holding its own away from human
settlements; the South American rain forest is still very large, and
the tortoises have a distinct tendency to remain more or less hid
den in very shady areas, avoiding open clearings and wide trails
where they are liable to be caught.
The red-foot tortoise, Geochelone carbonaria, was described
as a separate species by Spix as long ago as 1824, and the older
museum specimens of the two species of northern South American
tortoises usually bear labels making clear the distinction between
the two (though in the older literature denticulata is usually called
tabulata). However, recognition of the difference between these
two really quite separable species fell into disuse for some unac
countable reason until 1960, when Ernest Williams (Breviora,
M.C.Z., No. 120: 1-13) re-examined numerous museum specimens
and re-asserted the distinctness of the two forms.
Geochelone carbonaria is a smaller species than G. denticu
lata, adults usually being around a foot in length, and the record
length, as far as I know, being represented by a 17.75” shell in the
American Museum of Natural History. In contrast to denticulata,
males get a little larger than females. The coloration in typical
specimens is quite distinct from that of denticulata, the carapace
being black (hence the specific name), with a small, distinct yel
low area on each of the scutes; even in large specimens these yel
low markings do not extend much beyond that part of the scute
which corresponds to the size of the scute at hatching. The plastron
is relatively unpigmfented, but there may be some reddish tint and
vague dark markings along the areas of more recent growth, while
in many adult males the plastral concavity may be pigmented.
The skin is black, with striking bright yellow markings on the head
and lower jaw, including a strong horizontal bar behind the eye.
Many of the scales on the limbs and tail are bright scarlet in color.
Half-grown carbonaria have parallel-sided shells, but mature spe
cimens have a very distinctive incurving of the sides, which gives
326
The yellow-footed tortoise, Geochelone denticulata. Photo taken by Harald

Schultz in Brazil.
the animal a well-defined “waist.” The plastron is also concave in

adults; both lateral and ventral concavities are most strongly de
veloped in males.
The gular scutes of denticulata and of carbonaria are differ
ent; in denticulata they are normally longitudinally divided in dor
sal aspect, giving the appearance from above of four gular scutes.
Also the gulars extend further back along the midline in carbo
naria, so that they encroach upon the entoplastron and shorten the
interhumeral seam to the point that it is usually shorter than the
interfemoral seam. In denticulata the gulars do not encroach upon
the entoplastron, and the interhumeral seam is longer than the
interfemoral. There are also differences in the head scalation of
the two species; denticulata has elongate prefrontal scales and a
fragmented frontal scale. Carbonaria has short prefrontals and,
usually, a large, intact frontal. Denticulata has very small ingui
nal scutes, barely visible in ventral view, while carbonaria has
much larger inguinals, conspicuous in ventral view. The anterior-
most marginal scutes in very young denticulata show distinct den-
ticulations (hence the name of the species), while this region in
young carbonaria is almost smooth.
Specimens of carbonaria from west of the Andes appear
rather different from the typical coloration described above; the
327
www.Ebook777.com
Impressed tortoise, Geochelone impressa, a rare species from Southeast
Asia. Photo by Larry Lantz.
The Burmese brown tortoise, Geochelone emys.

Travancora tortoise, Geochelone travancorica, on the left and the elongated

tortoise, Geoochelone elongata, on the right. These two species from the In
dian subcontinent are closely related.
329
www.Ebook777.com
carapace is greyish or brownish rather than black, and the light
limb scales are yellowish or slightly orange, but not scarlet. This
coloration is seen in specimens from the Choco of Colombia, from
Panama (Darien and San Bias Provinces), and from Old Provi
dence Island. Many specimens from Colombia also show yellowish
rather than red scales on the limbs, but there is usually some con
trast between the color of the head scales and of the limb scales,
while there is no contrast in denticulata. In Ceara, Brazil, where
denticulata is absent, carbonaria consistently reaches a very large
size (16” or more), and both head and limb markings may vary
from yellow through orange to red, with or without contrast be
tween the two areas.
In Panama, carbonaria is reported to be a forest living ani
mal. In Surinam, however, the animal is more typical of savannas.
Marinus Hoogmoed informs me that in Surinam only carbonaria is
found in the savanna areas, while in the forested regions denticu
lata is dominant but a low density of carbonaria is also present.
The maintenance of at least some population in the forest areas is
necessary, since the savanna lands in the Guianas are frequently
surrounded by forest, and it is thought by some that they are en
tirely of human origin. The forest-living carbonaria individuals
thus provide a nucleus for the invasion of new savanna lands that
open up as a result of man’s ‘'slash-and-bum” agricultural
methods.
Medem reports that in Colombia carbonaria lays 5 to 15 eggs
from July to September. Local people in Panama informed Auffen-
berg that the eggs are not buried but are deposited in the leaf litter
on the forest floor. On the other hand, I have witnessed excavation
of a typical tortoise nest by a carbonaria in Guyana. Legler esti
mated from dissection of a series of Panamanian females that “8
to 13 eggs per clutch might be logically expected.”
The red-foot tortoise is just as esteemed as the yellow-foot as a
source of food; Legler reports that in Panama tortoise meat is
more popular among the San Bias Indians than any other wild
meat.
Auffenberg recently described Geochelone hestema, a fossil
tortoise from the Miocene of Colombia, which appears to be mor
phologically intermediate between denticulata and carbonaria
and which may be ancestral to these two species. He postulated
that the separation of denticulata and carbonaria may have occur
red with the development of savanna and grassland peripherally
to the Amazon basin in the Pliocene or Pleistocene. The two spe
cies may have become distinct in isolation as one became special
ized for forest life and the other for savanna life; but the two occur
sympatrically in many places today, and it is instructive to con
sider how the two species avoid interbreeding.
330
I remember an incident which occurred several years ago

when I was making inquiries about tortoises in the food market in
Surinam. A stout Surinaamer, of uncertain and complex racial
background, picked up an adult male and female denticulata from
a box behind his counter and, indicating-each animal in turn, ex
plained “dis one man; dis one vrouw; ” then, rattling the reluctant
and totally retracted animals together in an approximation of the
copulatory position, he went on “dis how dey fock, man.” Basic
ally, he was correct, but he left out the most interesting part, the
courtship behavior and sex and species recognition techniques
shown by this species and by G. carbonaria. These aspects have
now been studied by Walter Auffenberg (1965: Copeia 3, 335-342).
Although Auffenberg’s tortoises unfortunately did not come from
an area where the species are sympatric (carbonaria from Bar
buda; denticulata from Leticia, Colombia), his results are edify
ing. It appears that a male tortoise identifies males of its own spe
cies by eliciting a characteristic head movement; if this move
ment is not forthcoming, the male walks round to the back of the
other tortoise and sniffs at the cloacal region. A female of its own
species is immediately recognized, and copulation follows either
straight away or after a variable period of biting at the legs (pre
sumably a method of immobilizing an unwilling female).
The actual head movements of denticulata are as follows: the
tortoise usually challenges another from a side-by-side position by
means of sudden movements of the head from the normal, frontal
position to one side—usually the side on which the other tortoise is
situated—followed by a slightly slower return to the mid-position.
The other tortoise, if a male of the same species, will normally
respond with a similar movement. The head movement of carbo
naria, on the other hand, instead of being a single sweeping move
ment to one side with similar return movement, consists of a series
of jerks away from, and back to, mid-position. This is followed by a
similar series of movements to the opposite side. However, in car
bonaria the response is somewhat more specific than in denticu
lata-, the movements not only have to be as described above, but
the head (or artificial head, under experimental conditions) has to
have the usual vivid coloration typical of the species.
When copulating, male carbonaria emit a strange “clucking”
sound, amazingly reminiscent of that of a hen; it consists of a
series of fairly slow “clucks” of moderate pitch, followed by sever
al more of rapidly rising pitch and at decreasing intervals. In the
Crandon Park Zoo, I once saw two juvenile carbonaria, each only
about five inches long, going through the entire clucking and
mounting routine.
Until recently, it was customary to refer all other South
American tortoises to a single species, Geochelone chilensis, the
331
www.Ebook777.com
The Chaco tortoise, Geochelone chilensis, from La Posta, Cordoba, Argen
tina.
An excellently colored specimen of the red-footed tortoise, Geochelone car-

bonaria.
332
Nesting female Geochelone elephantopus becki, a giant tortoise from the

northernmost volcano of Isabela Island, Galapagos.
Adult male giant tortoise, Geochelone elephantopus porteri, from Santa Cruz
Island, Galapagos.
www.Ebook777.com
Chaco tortoise. This was thought to be a small form, not exceeding
a shell length of 11.1”, with most adults about eight inches long.
However, Marcos Freiberg has recently published descriptions of
two new species, G. donosobarrosi and G. petersi, based on speci
mens which would formerly have been identified as G. chilensis.
Freiberg has also demonstrated that even chilensis in the narrow
sense may reach a shell length of 36 cm, or about 14”, while the
relatively giant donosobarrosi may get even larger than that—a
specimen kept alive by Sr. Manuel Lucio Alcuaz, of Echeverry in
Buenos Aires Province, Argentina, has a straight length of 433 mm
or about 17”. Some question has been raised as to whether Frei
berg’s new species are valid, however, and Walter Auffenberg,
who is familiar with these tortoises in the field, informs me that he
believes donosobarrosi to be a subspecies of chilensis and petersi
to be based on male specimens of normal chilensis, the supposed
specific characters being involved with sexual dimorphism.
Provisionally accepting Auffenberg’s interpretation, G. chi
lensis has an essentially subtropical distribution in Paraguay,
northern and central Argentina, and probably also southern
Bolivia. Old records from Uruguay, Chile, and Buenos Aires,
Argentina are now considered invalid. The name chilensis is of
course based on a mislabeled specimen; when this mistake was
discovered Sclater immediately proposed the more appropriate
name “Testudo argentina. ” However, Gray’s introduction of the
name chilensis antedated Sclater’s publication by a few months, so
chilensis must stand despite the error on which it is based.
G. chilensis chilensis is a medium-sized tortoise with a low,
broadly oval carapace with the shell margin frequently strongly
serrate. The plastron is broad and the epiplastral lip is well devel
oped. The head is relatively large; the head scalation consists of
several large, elongated, rectangular scales in contact with one
large triangular (or two smaller rectangular) prefrontal scale,
with or without two smaller scales on either side. The frontal scale
is large, sometimes variously divided, lying behind and between
the eyes, and separated from the more anterior scales described
above by a row of small scales. The scales on the limbs are large,
pointed, and overlapping. There is a patch of enlarged scales on
the thigh region, one or more of which may be enlarged into a
small spur.
The Chaco tortoise is an overall ochre-yellow or yellow-brown
in color, often, especially in younger specimens, with somewhat
darker scute edges. This coloration appears to have definite cryp
tic value in the natural habitat, and specimens from dry, sandy
areas have lighter coloration than those from areas where the sub
strate is wetter and thus darker. The habitat of the Chaco tortoise
embraces a wide band of arid lowland bounded on the north by
334
rainforest, on the west by the Andes, and on the south by the cold
wastelands of Patagonia. The diet is reported to consist of fruits,
cactus, and grass. Nesting takes place in the southern summer,
mainly in January, but the season extends from November to
early March. One to four eggs are laid at a time (up to six, accord
ing to Freiberg); the eggs, as in many small tortoises, are
markedly elongate, measuring about 1.8” x 1.4”. Hatching may
take as long as a year under natural conditions.
Some sexual dimorphism is shown by the Chaco tortoise; the
males are reportedly smaller, with more serrate marginals,
longer and more divergent epiplastra, more acute and larger
xiphiplastral projections, and proportionately larger heads and
flatter shells. Females have thinner shells than males, often with
actual fenestrations, as do females of some of the Galapagos tor
toises. In general appearance the Chaco tortoise is amazingly
similar to the North American species Gopherus agassizi and G.
berlandieri; this reflects parallel evolution in a very similar habi
tat. The species may be immediately distinguished from Gopherus
by the absence of the nuchal scute.
At night Geochelone chilensis retires to a “pallet” or shallow
burrow very similar to that of Gopherus berlandieri. During the
cool season a deeper pallet is used, and the tortoise only comes out
on the warmest days. Another similarity to the Texas tortoise is
the high population density that is reached in suitable habitat; Auf-
fenberg (to whom I am indebted for much of my information about
this species) found a population density of 15-20 tortoises per acre
near La Posta, Cordoba, Argentina. Freiberg found them especial
ly common in Cordoba also, as well as in Santiago del Estero.
Adult Chaco tortoises seem to have no major natural preda
tors; however they are exported to Santiago, Chile for food, and
the young are sent to Buenos Aires and Mendoza for the pet trade.
The species is protected by law, but violations are frequent, and
Freiberg describes G. chilensis as “an endangered species.”
The subspecies donosobarrosi is found at the southern extreme
of the range of the species, the type specimens being from San
Antonio, Rio Negro; Colonia, near Rio Colorado, La Pampa; Car
men de Patagones; and General Acha, La Pampa (all in Argen
tina). Apart from its larger size, the subspecies is differentiated by
the darker shell, with black-centered scutes, the width of the pos
terior plastral lobe less than the width of the bridge, and the
shorter interhumeral seam (equal to the interabdominal seam).
The eggs, according to Freiberg, measure 38 x 49 mm.
The subgenus Chelonoidis also includes the famous giant tor
toises of the Galapagos Islands, located on the Equator in the East
Pacific, about six hundred miles west of Ecuador, and named after
their chelonian inhabitants. There were originally about fourteen
335
www.Ebook777.com
Giant tortoise, Geochelone elephantopus microphyes, from Volcan Darwin,
Isabela Island, Galapagos.
Three large male giant tortoises, Geochelone elephantopus vandenburghi, in

the caldera of Volcan Alcedo, Isabela Island, Galapagos.
Young giant tortoise, Geochelone elephantopus guntheri, from Sierra Negra,

Giant tortoise, Geochelone elephantopus elephantopus, from Cerro Azul,

www.Ebook777.com
isolated populations of Galapagos tortoises, each separated from
its neighbors either by the ocean or, in the case of the four or five
races of Albemarle Island, by stretches of impassable fresh lava.
These populations were each considered to constitute separate
species by many early writers, but now they are all regarded as
subspecies of Geochelone elephantopus. On the larger islands the
tortoises may reach weights of 500 pounds and more, and carapace
lengths of four feet, but apart from their enormous size they show
relatively few modifications from the normal mainland tortoise
condition. These few differences include the absence of bony cores
in the scales on the front faces of the forelimbs and the relatively
long neck and small head. On the other hand, the tortoises on the
smaller and drier islands appear to be runted and somewhat mal
formed; it is only on closer investigation that one realizes that
these “malformations” tend to be relatively constant within the
population of a particular island, and moreover that they are prob
ably highly adaptive traits.
These differences shown by the small island tortoises may be
summarized as follows: i) the animals as a whole are smaller; fe
males average sixty pounds or less and males average not more
than 120 pounds; ii) the carapace shows a so-called “saddleback”
shape, being distinctly elevated above the neck instead of curving
downward as in other tortoises; iii) the carapace is strongly flared
or even reverted above the hind feet; iv) the limbs are longer and
thinner. The neck also appears to be longer, but this is probably
simply because the saddle-fronted shell allows the entire neck to
be raised almost vertically, while in the dome-shelled tortoises the
basal part of the neck is necessarily bent downward even when the
tortoise is reaching up as high as it can. The head and neck are
more slender in most saddleback races than in the dome-shelled
ones.
To interpret the meaning of these differences between the
dome-shelled and the saddle-backed Galapagos tortoises, we
should realize that the ancestral Galapagos tortoise certainly must
have had the domed shapes'; since the open front of a saddleback
shell would offer no defense against large predators. However, in
the isolation and security of the Galapagos Islands, certain popula
tions evolved saddled shells with correlated differences in the ex
tremities.
Under primordial conditions, Galapagos tortoises reached
phenomenally high population densities. Nevertheless there must
ultimately have been some sort of limiting factors on the popula
tion size, since they did not go on expanding to infinity. It appears
probable that on the smaller, lower, and more arid islands, such as
Duncan and Hood, food availability would be strongly limiting,
especially during prolonged rainless periods. We can see that a
338
Large number of Geochelone elephantopus vandenburghi at a waterhole on

Volcan Acedo, Isabela.
tortoise that could extend its head a little higher than its neighbors
would be able to reach vegetation that was out of reach of the
others and thus would be at a distinct advantage. This appears to
be what has happened. A saddleback tortoise can reach upward to
an amazing height and has essentially become a browser as well
as a grazer. In this connection it is interesting to note that one of
the principal sources of water on the arid small islands in the Gala
pagos, or in the arid parts of the larger islands, is the fleshy pads of
the Opuntia cactus. In continental areas, and on those Galapagos
Islands (such as Tower and Bindloe) which have never had giant
tortoises, the cactus assumes so-called recumbent posture,
spreading out close to the ground. On the other hand, on the tor
toise islands the cacti have evolved an arborescent form; the
young plants, which are exceedingly spinose, may put on a dozen
or more pads absolutely vertically before any branching occurs,
and then they proceed to form a trunk, covered with inedible
brown bark, out of these vertical pads. The tortoises thus can only
eat those pads which grow down again toward the ground, and the
plant is in no danger of being destroyed when these pads are eaten.
339
www.Ebook777.com
Large male James Island giant tortoise, Geochelone elephantopus darwini,
from the Galapagos Islands.
Adult male giant tortoise from Hood Island, Geochelone elephantopus

a race on the brink of extinction but now recovering.
hoodensis,
341
www.Ebook777.com
Galapagos tortoise, Geochelone elephantopus ephippium, from Duncan Island.
The theory assumes that some factor other than food shortage
limited the tortoise populations on the more fertile islands. I be
lieve that the limiting factor here was availability of nesting
space. The tortoises form huge populations in the highlands, where
rainfall is high and vegetation luxuriant, but the eggs have to be
deposited in the hot lowlands if they are to have a good chance of
hatching. However, the lowlands of the islands are for the most
part composed of sheets or blocks of lava, and the areas of fine soil
of adequate depth for nesting, which also are accessible from the
highlands, may be very limited. Thus, when the tortoise popula
tions become very high, the relatively few suitable nesting areas
will be overcrowded and the female tortoises will destroy each
other’s eggs in their efforts to bury their own. There will thus be
lower overall recruitment to the population until the population of
adults has dropped. However, it should be mentioned that even on
the small dry island of Duncan, where the tortoise population is
very reduced from former times, some nest interference of this
type still takes place.
It is also possible that the cut-back shell, flared hind shell mar
gins, and long legs of the saddleback tortoises assist them in tra
versing the often extremely rugged terrain of their native islands.
The open shell front, with much exposed skin, may also serve a
thermoregulatory function, since the smaller Galapagos Islands
are often exceedingly hot. Nevertheless, one might ask why it is
that, among the saddleback tortoises, it is the adult males that
show the condition in its extreme form, when presumably the fe
males have just as much need, if not more need, for the additional
food that greater height would bring within their reach. The
answer is probably that carapace saddling grows allometrically—
that is, it is absent in the hatchlings and appears gradually as the
tortoise grows. Since males grow much larger than females, they
show the condition in a more developed form. Males and females
of equal size are probably about equally saddlebacked. It is likely
that the saddlebacked-shape has failed to develop on the larger,
more humid islands not only because it is not necessary, but also
because the high front of a saddleback tortoise offers a definite im
pediment when the animal is pushing its way through dense vege
tation. A dome-shelled tortoise, on the other hand, can walk
through such vegetation because the shell front is able to push
branches and vines upward and out of the way.
The sexual dimorphism of Galapagos tortoises, with males so
much larger than females, is worth further discussion, since it is
the reverse of the typical condition among mainland tortoises.
Under mainland conditions, where it is not unusual for two or more
tortoise species to occur sympatrically, male tortoises of many
species have evolved complicated rituals for identifying females
342
of their own kind and “persuading” them to remain still so that

they can be mounted. Other things being equal, it is advantageous
for the female to be large, since this enables her to carry more, or
larger eggs. However, in the Galapagos, where the total egg
volume is very small by comparison with the volume of the female
tortoise, there is no advantage in the female being larger than the
male; and the males, instead of evolving complex “persuasive”
techniques to immobilize the female, court the females simply by
battering them with the front part of their shells, and then bearing
them to the ground by sheer superior weight before mounting.
Copulation itself among the Galapagos tortoises may occur in
almost any month of the year, though it does show seasonal peaks.
When actually in the copulatory position, the male tortoise emits
deep bellowing groans which can be heard at a great distance.
Galapagos tortoises may batter and bite each other at other
times than when mating, and fights often involve individuals of the
same sex. The tortoises have a very characteristic style of ritualis
tic combat; they face each other and each raises his head as high
as he can in order to deliver a downward bite on his opponent’s
head. The fight immediately terminates when one tortoise has
demonstrated that it has the higher reach, and the loser usually
makes off at high speed.
Galapagos tortoises are capable of eating almost any green
vegetation that they may encounter, though one or two species
(e.g. Maytenus octagona) are avoided. On some islands, the tor
toises feed upon large quantities of the “manzanillo” (Hippomane
mancinella), a small green fruit that is highly poisonous to most
organisms. When the opportunity presents itself they spend much
of their time partially submerged in water. This may be not only
for thermoregulatory purposes, but also to lessen attacks by mos
quitoes and ticks. At night they often dig into soft ground or vegeta
tion so that only the hind part of the animal remains exposed. It
has been suggested by some authors (e.g. J.R. Hendrickson) that
wallowing in waterholes assists in some way with respiration; but
this argument is not now considered valid, for reasons sum
marized by Gans and Hughes (1967: J. Exp. Biol. 47: 1-20).
At the present time, Indefatigable Island is the place where
most Galapagos visitors are recommended to go if they wish to see
the giant tortoises in the wild state; tortoises are still numerous in
the “tortoise reserve” that has been established in the southwes
tern part of the island, and they may be visited by making a five-
hour hike along rough trails from the settlement at Academy Bay.
Moreover, the females descend quite low in the breeding season,
and may be seen after only a short walk from the south-west coast.
However, the tortoises of this island are barely mentioned in the
eighteenth century whalers’ accounts, and Commander Cookson
343
www.Ebook777.com
The only surviving individual of the giant tortoise from Abingdon Island,
Geochelone elephantopus abingdoni, an adult male discovered in 1971.
Giant tortoise, Geochelone elephantopus chathamensis, from Chatham

Island in the Galapagos.
The only known specimen of

the giant Fernandina Island
tortoise, Geochelone elephan
topus phantastica, a
subspecies now possibly ex
tinct.
345
www.Ebook777.com
wrote in 1875 that the tortoises of Indefatigable were so reduced
that they were no longer hunted, the few left being in the most in
accessible parts of the island; he was assured that a search of a
fortnight might not result in finding a single individual. The report
of the California Academy of Sciences party, which sailed to the
Galapagos on the schooner Academy in 1905, to the effect that the
tortoises were still fairly common probably reflects simply that
earlier visitors had looked in the wrong places rather than that
their numbers had undergone a recent increase. The Indefatigable
tortoises were heavily exploited for their oil until at least the
1930’s, and even in the 1960’s large numbers of shells of tortoises
killed by man were to be found in the southwestern part of the
island.
The Indefatigable tortoises are currently known as Geoche-
lone elephantopus porteri. The names planiceps and nigrita are
both older than porteri, but the former is based only on a single
skull of unknown origin and the latter on two tortoises of unknown
origin and of too young an age to show any subspecific characters.
The name porteri, being based on an adult male definitely collec
ted on Indefatigable, is thus more satisfactory. This is a large
form, reaching a weight of over 500 lbs., and is very similar in
shape to the Central and South Albemarle races. However, the
shell is somewhat rounder and more highly domed, and there is no
sign of elevation or narrowing of the anterior margin. The overall
color is black, with a variable whitish to yellowish patch under the
throat of adults.
At the present time, the Ecuadorian National Park Service
personnel have marked 1420 Indefatigable tortoises, and the total
population is estimated to number around 2000-3000. Moderate
numbers of young exist, and most of the specimens one sees are
about half grown. Reproductive success was, however, diminished
for many years by the presence of feral pigs in the nesting areas.
Personnel of the Park Service have made an intensive effort to
eliminate these pigs indecent years and to protect the nests by con
structing corrals around them out of lava rocks. Peter Kramer re
ports that during the 1971-72 nesting season 115 nests with a total of
1,052 eggs were protected in this way, and that 706 young tortoises
successfully hatched. These figures give the average clutch as
9.16, but the maximum clutch size for Indefatigable is 17. There is
some suggestion that tortoises may lay more eggs in rainy years
than in dry ones, and certainly those races whose habitat is consis
tently wetter produce larger clutches than those from dry islands.
On Indefatigable, each female tortoise may lay twice in a season,
the first clutch often being larger than the second.
The Indefatigable tortoises nest in so-called “campos”—areas
of little vegetation with silty soil in the lowlands of the southwes-
346
The Galapagos Islands. I: Abingdon (Pinta); 2: Bindloe (Marchena);

3: Tower (Genovesa); 4: Chatham (San Cristobal); 5: Hood (Espanola);
6: Barrington (Santa Fe): 7: Charles (Floreana);7 8: N. Seymour;
9: S. Seymour (Baltra); 10: Indefatigable (Santa Cruz); 11: Duncan
(Pinzon); 12: Jervis (Rabida); 13: James (San Salvador;14: Bartholomew;
15: Albemarle (Isabela); 16: Narborough (Fernandina); 17: Crossman
(Los Hermanos); 18: Brattle (Tortuga); 19: Darwin (Culpepper);
20: Wolf (Wenman).
tern part of the island; these they reach by means of a system of

trails, trodden out by generations of tortoises, which extend back
into the upland feeding areas. These “campos” have been
described by Hendrickson (1965: Pacific Discovery XVIII, 5: 28-
36). Most are composed of a very fine-grained red-brown soil
which is thoroughly mixed with the deteriorating fragments of tor
toise eggshell, the product of centuries of nesting in the same
areas. The nesting tortoise chooses a spot which receives full sun
light and makes a nest cavity by urinating on the ground and dig
ging out a hole in the mud with her columnar hind feet. The final
cavity is about 7” to 12” deep and about 8” to 10” in diameter. Hen
drickson reported that the nest is closed by the tortoise lowering
her plastron over the cavity and excavated mud and sliding about
to produce a plastered cap nearly flush with the surrounding soil;
347
www.Ebook777.com
348
Margined tortoise, Testudo marginata, from Greece.
Afghan tortoise, Testudo horsfieldi, a small, cold-hardy species from

Pakistan, Afghanistan, eastern Iran, and southern USSR.
www.Ebook777.com
Outline of shell of Geochelone
elephantopus porteri.
but Craig MacFarland did not witness this behavior in any of fif
teen nestings. The muddy area normally dries to an almost brick-
hard consistency in the tropical sun, which is probably advanta
geous in preventing the eggs from dehydrating or being broken if
another tortoise walks over the spot.
It appears possible that at one time tortoises were found over a
much wider area of southern Indefatigable. There is now an exten
sive farming community in the south-central part of the island east
of the tortoise reserve, and only occasional stray tortoises are now
found in this altered habitat. A few dozen tortoises persist in an un
disturbed area east of the farm lands, but there is no recent infor
mation on this population, and they may now be so dispersed that
breeding has ceased.
There is one other area on Indefatigable where tortoises have
been seen in recent years. This is in the northwestern part of the
island at least six miles from the nearest part of the tortoise re
serve, near an isolated coastal mountain known as Saddleback
Hill. A large male tortoise found here by the Angermeyer brothers
of Academy Bay in 1962 was quite different in shape from normal
porteri, having an extreme saddleback shape reminiscent of
(though not identical with) that of the Abingdon tortoise (abing-
doni). David Snow, then Director of the Darwin Station, visited the
area with the Angermeyers in 1963, and not only located and photo
graphed the big saddleback, but also found six juvenile tortoises in
the same area, and a seventh dead juvenile. There has been no
subsequent information on this population with the exception that I
paid a brief visit to Saddleback Hill in early 1972, and found a 13”
tortoise, which judging by its size was hatched since Snow visited
the area. It thus appears that there is a small breeding population,
but how it got there, and from where (assuming indeed that it is
introduced and not natural), remain unknown.
Albemarle Island, the largest of the Galapagos, is an irregu
larly-shaped island about 75 miles long composed essentially of
five volcanoes, each with a caldera several miles in diameter, with
a sixth isolated massif at the northwestern tip known as Cape'
Berkeley or Volcan Ecuador. A separate subspecies of tortoise has
been described for each of the main volcanoes, and these indeed
appear to be as distinguishable as any on the isolated islands, with
350
the exception of the two southernmost forms which at least poten

tially have genetic contact across the fertile lowlands between the
two craters. The three northern populations are all separated from
each other by miles of shadeless and impassable lava.
351
www.Ebook777.com
Psammobates oculifer, a dwarf

South African species commonly
known as the serrated tortoise.
South American geometric tortoise, Psammobates geometricus, one of the

rarest living tortoises. Photo by James Juvik.
353
www.Ebook777.com
Volcan Wolf is the northernmost volcano of Albemarle and is
the highest mountain in Galapagos. It is a relatively sterile area
with exceedingly precipitous terrain, and it is probable that the
tortoises here never assumed the population densities originally
present on South Albemarle. We do not know the extent to which
tortoises on this volcano were despoiled by whalers (if indeed they
were at all), since most logbook records simply mention “Albe
marle” without specifying which part. The Academy party found
seven tortoises in seven days near Bank’s Bay, on the west side of
Volcan Wolf, and at the present time the population appears to be
moderately numerous, widespread on both the northern and wes
tern slopes of the mountain, and reproducing well—most of the tor
toises one finds are young or half grown. Craig MacFarland esti
mates that over a thousand tortoises may still survive on this
mountain, but this is only an educated guess as none of the tor
toises have been marked. I found seven individuals in a few hours
near Bank’s Bay in 1971, and three more near Punta Albemarle in
1972. The tortoises sometimes are found very close to the sea, and
this accessibility makes them liable to depredation even today by
poachers collecting for unscrupulous zoos and private collectors.
Tortoises on Volcan Wolf show a bewildering variety of cara
pace shapes; some quite large specimens are dome-shaped, others
have very flat shells, and still others, even of small size, may be
strongly saddlebacked. An amazing photograph taken near Bank’s
Bay several years ago by Daniel Weber, and published in Roger
Perry’s book The Galapagos Islands, shows a tortoise whose shell
is fully as saddlebacked as a mature male from Hood Island, sit
ting just beside a typically dome-shaped animal. The reasons for
this variability are unclear; possibly the variety one sees is the
result of a relatively recent coming together of thitherto isolated
saddlebacked and dome-shelled populations.
Few fully mature tortoises from this mountain have been seen
in recent years, but the largest male collected by the Academy
part in 1906 was 41.5” long, and the type specimen in the Tring
Museum is 40.75” inilength. Those individuals which are saddle
backed are not narrowed anteriorly as are most of the saddle
backed races, but instead many have a broad and flaring anterior
shell opening. Even relatively small specimens may have quite
smooth scutes, especially on the plastron, where the growth annuli
are presumably worn smooth by frequent contact with very rough,
rocky ground.
The breeding habits of the Volcan Wolf tortoise are unknown.
Reproduction appears to be fairly successful, even though both
black rats and feral cats are now found on this mountain—the
effects of these animals on the tortoises are unknown at present.
The type specimen of Geochelone elephantopus becki was sup
posedly collected at Cape Berkeley, which is in fact isolated from
354
Volcan Wolf by several miles of fresh lava. Recent visitors to Cape

Berkeley have found no signs of tortoises there, hut it remains just
possible that tortoises live on an almost inaccessible plateau about
two-thirds of the way up the northern side of the mountain. If this
area indeed proves to have a tortoise population, then the popula
tion on Volcan Wolf will have to be re-named.
Volcan Darwin, just south of Volcan Wolf, is the home of the
subspecies Geochelone elephantopus microphyes. The tortoises
are found principally in the highlands to the south and west of the
main crater, but females descend very low on the western slopes
during the breeding season. There is an excellent protected
anchorage at Tagus Cove on the west coast of Volcan Darwin, so
this population provided an important source of supply of live tor
toises to the nineteenth century whalers. The Academy party in
1906 found that the preferential removal of the more accessible
female tortoises by whalers had caused a considerable imbalance
in the sex ratio—they found thirteen males and only one very
young female. At the present time it is estimated that there are be
tween 500 and 1000 tortoises on Volcan Darwin, but this estimate is
also only an educated guess, and only 65 tortoises from this popula
tion have been marked. The population appears to be breeding
successfully despite the presence of cats and rats, but as on Volcan
Wolf more information is needed. The sexual imbalance noticed at
the beginning of this century seems to have evened out, and a high
proportion of the tortoises one finds today are immature. Not only
do the breeding females migrate up and down the side of the moun
tain, but the population as a whole tends to move to lower areas to
the west during rainy periods and to move back up to the moister
highlands during very dry weather. When I climbed Volcan Dar
win from Tagus Cove in January 1971, conditions were very dry
and I saw no tortoises in the lowlands, only one young specimen
half way up the mountain, and no adults until I was quite close to
the crater rim. However, when I revisited the area in March 1972,
the weather was wetter than it had been for many years, and I
counted seventeen tortoises, of all ages and both sexes, in a low
land vegetated part of the valley between Volcan Darwin and
Tagus Cove.
The Volcan Darwin tortoise is a medium-size race with a very
thin and rather flattened shell. The front of the carapace is not
elevated, but in mature males the anterior carapace margin is ex
tended forward to form a horizontal flange. In general appearance
this race is rather similar to that of James Island, but it does not
reach such a large size—the largest microphyes on record had a
carapace length of 40.5 inches. The chin and throat are not marked
with yellow or white in this race.
It is known that microphyes nests in lowland areas at the wes
tern foot of Volcan Darwin and on the slopes just above. The
355
www.Ebook777.com
Psammobates tentorius verroxii, a race of the South African tent tortoise
distinguished by its relatively flat dorsal scutes. Photo by John Visser.
A pair of pancake tortoises, Malacochersus tornieri, from Kenya. This ex

traordinary species has a soft, flattened shell and takes refuge in rocky
crevices when disturbed.
Two views of the South African bowsprit tortoise, Chersine angulata.
357
www.Ebook777.com
average clutch size is probably about eight, based on six clutches
counted by Craig MacFarland.
Volcan Alcedo, to the south of Volcan Darwin, is the lowest of
the Albemarle volcanoes, the rim being 3650 feet above sea level.
The anchorages, at least on the east side of the mountain, are poor,
and the country is so open that no tortoises are found for several
miles from the sea, so that the whalers appear to have left this
colony alone. However, Galapagos settlers were in the habit of
camping on the side of the mountain in order to hunt tortoises for
their oil, and the Academy party in 1906 found about seventy old
skeletons in one such camp. They only managed to find one living
tortoise, however, and concluded that the natives had cleaned the
tortoises out very thoroughly.
It was not until about 1965 that Miguel Castro succeeded in
finding a route through the dense vegetation that prevented the
Academy party from reaching the actual rim and interior of the
caldera of Volcan Alcedo, and he reported a large tortoise popula
tion that subsequent visitors have estimated to number from 3000
to 5000 animals, and thus the largest in the archipelago today. Un
like the other Albemarle craters, the Alcedo caldera is fertile and
most of the tortoises on the mountain live and breed actually in the
crater and on the wet southeastern, southern, and southwestern
outer flanks. Paths up and down the inner slopes of the crater walls
allow the tortoises to come and go as they wish; and during the wet
season they may descend some way down the east side of the
mountain. Until about 1971, the tortoises were found in particularly
high concentrations in a muddy pool formed by the overflow from
a boiling geyser a third of the way down the inner slope of the
south-west wall of the caldera. However, the geyser has now dried
up, though it still emits steam. The overflow pool re-filled with
rainwater during 1972, which was a very wet year, and tortoises
were once again able to drink and submerge themselves there.
The Alcedo tortoises are dome-shelled, very similar to the In
defatigable tortoises in shape, but the shell is a little less rounded.
The larger scutes are moderately bossed, and there is a slight nar
rowing in the region of the anterior parts of the lower edges of the
first costal scutes. The maximum size of adult males is very great,
the biggest ones having carapace lengths of more than fifty inches.
Females are much smaller—around thirty inches long. The popu
lation is breeding quite well, there being large numbers of individ
uals 12” to 18” in length, though relatively few smaller than that;
on my four visits to this mountain I only found one really small tor
toise (about four inches long). Copulation is often observed and
probably takes place in almost all months of the year. I noticed a
great deal of copulatory activity in March 1973.
358
The outlook for the Alcedo tortoises appears good, although

cats and rats have been reported from this mountain and there are
numerous feral donkeys which have the habit of rolling in the tor
toise nesting areas, with potential damage to both eggs and hatch
lings. The donkeys probably also destroy occasional young tor
toises, up to 15” or so in length, by trampling them.
The Alcedo tortoises are normally called Geochelone elephan-
topus vandenburghi, but the legality of this name is somewhat
questionable. Despite the abundance of the race, there is only one
complete museum specimen in existence, since the Galapagos tor
toises have been completely protected since before this population
was rediscovered. The single specimen in the California Academy
of Sciences, a young mature female, was collected by the
Academy party in 1906. Van Denburgh declined to propose a for
mal scientific name for the race since he had no idea of the full
range of variation of the population. The name Testudo vanden
burghi was proposed by De Sola (1930: Copeia 3: 79-80) after he
had visited a tortoise population south of Perry Isthmus—i.e. on
the lower slopes of Sierra Negra and isolated by impassable lava
from the Alcedo population. Moreover De Sola neglected either to
designate a type specimen or to give any information on how his
new species could be distinguished from its relatives.
The huge block of land that makes up the southern half of Albe
marle Island was formerly inhabited by very large populations of
tortoises of great size. These populations persisted in nearly pri
mordial abundance until the latter half of the nineteenth century.
During that period settlements were established on the island and
the collection of tortoise oil became a fairly large-scale industry.
Beck (1903: Seventh Annual. Report New York Zoological Society,
reprint pp. 1-17) gave a graphic account of the pitiful and merci
less way in which the tortoises were being decimated on the fertile
southern slopes of Sierra Negra, the more eastern of the two vol
canoes that make up southern Albemarle. To quote Beck:
“Some evenings, three men coming from different
directions would each carry in his hand a small piece of tor-
toise meat, and a pound or so of fat in which to cook it. Of
each tortoise killed not over five pounds of meat would be
taken, the remainder being left for the wild pigs that
swarmed about. ”
Beck took a photograph of literally hundreds of skeletons of
tortoises which had been killed by oil hunters. This photograph is
reproduced in William Beebe’s book Galapagos: World’s End.
These excesses, combined with the introduction of feral pigs,
dogs, goats, cattle, rats, donkeys, and cats, caused the complete
extirpation of tortoises throughout the fertile uplands of Sierra
359
www.Ebook777.com
Madagascan spider tortoise, Pyxis arachnoides, a very small species with a
distinctive anterior plastral hinge.
360
Madagascan flat-tailed tortoise, Acinixys planicauda, with detail of tail

region.
www.Ebook777.com
Negra and most of the valley between Sierra and Cerro Azul. How
ever, tortoises still exist in some of the more arid areas of southern
Albemarle. De Sola in 1928 found an apparently thriving popula
tion of tortoises, including breeding adults, in a coastal area near
Cartago Bay, just south of Perry Isthmus. Well over 50 individuals
were collected, and many others were seen. Unfortunately there is
no recent information on the present status of this population, if
indeed it still survives. In the arid area east of the settlement of
Santo Tomas, between Cerro Grande and Cerro Ballena, there is
still a population of tortoises which is managing to breed quite
well—indeed almost all of the tortoises found here have been of im
mature size. Personnel of the Galapagos National Park Service
have now marked 178 tortoises from this population and estimate
the total to be between 200 and 300 animals. The nesting area
appears to be at a relatively low altitude near La Cazuela, down
toward Cerro Ballena, right on the edge of the plate lava area.
Tortoises are also found on the south coast of Albemarle, in the
region of Cape Rose and Cabo San Pedro. This area is about equi
distant from the craters of Sierra Negra and of Cerro Azul, and the
existence of tortoises here suggests that originally a single tortoise
population extended across the entire south-facing slopes of
southern Albemarle. Van Denburgh considered the Cape Rose tor
toises to be referable to the subspecies microphyes—i.e., to be
identifical to the Volcan Darwin population. This allocation makes
no sense zoogeographically, though it is true that the populations
are very similar morphologically.
Tortoises are also found in a lowland, very moist and fertile
area north of Iguana Cove, on the west coast of Cerro Azul, as well
as in the inland lowlands east of the Cerro Azul crater. The tor
toises range all the way up to the caldera rim. Nesting zones exist
north of Iguana Cove, west of Cabo San Pedro, and north and east
of the main crater, on the edges of the fresh lava areas. 203 Cerro
Azul tortoises have now been marked, and the population is esti
mated to number 400-600.
The tortoises of Sierra Negra are usually called Geochelone
elephantopus guntheri, and those of Cerro Azul, G.e. elephantopus
(formerly G.e. vicina). The two races, if they are distinct at all,
are very similar, the differences described being really visible
only in adult males. Both reach a large size and have dome-shaped
rather than saddlebacked shells. However, adult male guntheri
tend to be rather flat, broad-shelled animals, while adult males of
elephantopus have more elevated shells, with a slight elevation
also of the anterior margin of the carapace.
The South Albemarle tortoises have the potential of growing to
an enormous size. Although really big specimens are not found
today, Baur in 1891 found a tortoise with a straight carapace length
362
of 49.5 ” and with a skull 7.12” long, about one day’s hike inland
from the old Cobos settlement. A party from the schooner ‘Mary
Sachs’ in 1902 found what Beck described as the largest known
Galapagos tortoise near the rim of Sierra Negra, about 18 miles
from the shore; it required the efforts of twelve men to carry it the
last six miles of the distance back to the ship. However, even
though the tortoises of Cerro Azul appear to be almost as large as
those of Sierra Negra, the eggs of the Cerro Azul tortoises are
smaller than those of any other race in the Galapagos (nine Iguana
Cove clutches of elephantopus showed an egg diameter range of 59
to 65 mm).
Beck found fresh nests on Sierra Negra in early November.
Clutch size ranged from eight to seventeen, and the eggs were laid
in three layers, with soft friable earth between each layer, with a
hard plug of dried mud between the top layer and the surface. Jose
Villa, of the Galapagos Park Service, found that clutches on Cerro
Azul average nine eggs per clutch and that nesting took place from
July to October. Young tortoises on Cerro Azul are menaced not
only by feral pigs, but also by ferocious feral dogs which hunt in
packs and regularly kill tortoises up to about a foot in carapace
length, and occasionally much larger ones. However, fair numbers
of young survive in the inland nesting area east of the crater,
where feral predators appear to be scarce; but there are almost no
young surviving in all the other nesting areas.
In 1928, when it appeared that there was no chance of the Gala
pagos tortoises continuing to survive in the wild state, the New
York Zoological Society dispatched an expedition to bring back a
large number of tortoises for distribution among various potential
captive breeding centers. Over two hundred South Albemarle tor
toises were collected, nearly all in the 7 to 25 pound range. Groups
of these were deposited in zoos and other institutions in Honolulu,
Bermuda, California (San Diego), Texas (Houston and San Anto
nio), Louisiana (New Orleans), Arizona (Superior), and Florida
(Brighton, Opa-Locka, Key Biscayne, and Lignum Vitae Key).
The tortoises being immature, it was some years before reproduc
tion was expected. Some tortoises from this distribution are still
alive in most of these places, but quantitative reproduction has
been achieved only in Honolulu and San Diego, though a few hatch
lings have been produced in Bermuda and Key Biscayne. In recent
years, Jack Throp of the Honolulu Zoo has been doing exceedingly
well breeding Galapagos tortoises; at the present time, 191 eggs
have been incubated and 60 hatchlings produced. The survivorship
of the hatchlings has been over 90%.
It is possible that a South Albemarle tortoise holds the record
for rapid growth. A 29 lb. specimen received by Edmund Heller at
Riverside, California, in 1899 had achieved a weight of 350 lbs. by
363
www.Ebook777.com
364
Florida snapping turtle (larger specimen), Chelydra serpentina osceola, with an unusual reddish phase of the common snapping tur
tle, C. s. serpentina.
Head of an Ecuadorian snapping turtle, Chelydra serpentina acutirostris.
A Mexican snapping turtle, Chelydra serpentina rossignoni, from Veracruz.
www.Ebook777.com
1906. It weighed 415 lbs. in 1913. It was kept in a muddy pen, 60 ft.
square, together with a cow; there was no opportunity for grazing,
but abundant food was provided. The tortoise died of enteritis in
1914; this is a common cause of death among tortoises under cap
tive conditions when they are exposed to very warm days and cold
nights; they eat a lot during the heat of the day, but are unable to
digest it when they cool down at night, and the mass of undigested
food sets up intestinal infection.
When Charles Darwin visited James Island in 1835, he found a
party of Spaniards encamped some six miles from the coast, en
gaged fulltime in hunting tortoises. While visiting them, Darwin
lived entirely upon tortoise meat, pronouncing that:
. . the breast-plate roasted (as the Gaucho do came
con cuero), with the flesh on it, is very good; and the young
tortoises make excellent soup; but otherwise the meat to my
taste is indifferent. ”
Townsend, in his study of whalers’ logbooks, found that large
numbers of tortoises were removed from James in the 1830’s, but
he could find no record of collection of tortoises from this island
later than 1845, in which year twenty were reported removed. It is
probable that by this time the tortoises in the more accessible
coastal areas had been drastically reduced, and the few that re
mained had migrated inland as the coastal lowlands were gradual
ly transformed into deserts by the introduced goats. These goats
had increased greatly upon the island, and at the present time
probably number at least 50,000 animals. The Academy party in
1906 made excursions into the interior of James from the eastern
end, but only found five tortoises, and Van Denburgh pronounced
them “very near extinction.”
Personnel of the Galapagos National Park Service, under the
leadership of Jose Villa, have recently cut a trail into the thitherto
virtually inpenetrable moist highlands in west-central James
Island, starting from the southern end of James Bay. Their find
ings were better than one dared expect, to the extent that they
have now marked 399 tortoises, out of an estimated total of 500 to
700 on the island. However, the sex ratio is distinctly unbalanced in
favor of males, and only about ten really young animals were
found, and very few of less than fully adult size. Villa estimates
that reproductive success diminished sharply about fifty years
ago, and stopped completely about thirty years ago. This repro
ductive failure can be directly traced to the presence of thousands
of feral pigs on the island, which tend to concentrate in the nesting
areas and destroy the eggs and hatchling tortoises with almost one
hundred percent efficiency. Jose Villa and two wardens shot about
1500 pigs during about four months on James Island in 1970, but
Villa estimates that at least twice that number remain, and their
366
Outline of shell of Geochelone e. dar-

wini.
reproductive potential is very high. In the meantime, eggs and

hatchling tortoises are being transferred to the Charles Darwin
Research Station, where they are maintained with a view to even
tual release of the tortoises back on James Island. By August 1972,
71 baby James Island tortoises had been hatched and were being
raised at the Station.
Villa found nearly all of the tortoises on James to be at alti
tudes of between 250 and 700 meters, though one or two were below
200 meters. The tortoises reach a very large size; the largest found
had a carapace length (over the curve) of 58.8”, which would cor
respond to a straight-line length of about 47”. The carapace is only
moderately domed, and the front margin is slightly elevated; the
race could be considered an intermediate form between the dome-
shelled tortoises and the saddleback races, rather similar to
microphyes of Volcan Darwin but larger and with a thicker shell.
The James Island tortoises, which are known as Geochelone
elephantopus darwini, are found in four distinct, though intercon
nected, zones in west-central James Island, and each of these
zones has a corresponding nesting area at relatively low altitude.
Most nesting takes place from August to October, and the eggs
typically number between 4 and 7 in a nest, with a maximum of 10.
Despite the tremendous abundance of feral goats on James Island,
there is no shortage of food for the adult tortoises in the highlands,
where there is sufficient moisture to sustain a vast area of lush
grassland known as the “pampas.” I visited this area in the com
pany of Mr. Leonardo Apolo, the only inhabitant of the island, in
January 1971. We covered a great distance in the cool, foggy up
lands, and found about twenty tortoises, almost all isolated large
males, for the most part sitting motionless in the pools of water left
by the recent rains. Some of these pools, such as “Poza Tragica”
and “Poza Felicidad,” appear to be permanent, and tortoises may
always be found in their immediate vicinity. I found one tortoise
(an adult female) during a day’s hike starting from Sullivan Bay,
at the east end of the island, in 1973.
Duncan Island, although only about three miles in diameter,
still has a small population of tortoises, of a distinctive saddleback
form. This island was relatively little despoiled by the nineteenth
367
www.Ebook777.com
Young alligator snapping turtle, Macroclemys temmincki, showing the worm
like “lure” on the tongue.
OPPOSITE: Two specimens of the Florida mud turtle, Kinosternon

subrubrum a subspecies with a smaller plastron, narrower
steindachneri,
bridges, and more aggressive disposition than the eastern mud turtle.
368
369
www.Ebook777.com
century whalers, and Townsend only found four logbook records of
whalers having removed tortoises from Duncan. These records
showed that fifty tortoises were removed from the island in 1848,
131 in 1850, 17 in 1855, and 208 in 1863. This relatively infrequent
visitation by whalers was probably partly because the island does
not have very satisfactory anchorages for large vessels, and
partly because the island is rather central in location, while
whalers normally filled up with tortoises at the first tortoise island
they came to—usually Abingdon if coming from the north, Hood or
Chatham if coming from the south.
After the whaling period, quite large numbers of tortoises
were removed from Duncan by scientific and semi-scientific par
ties. A number were collected on Duncan by the “Albatross”
expedition in 1888, and eight more were removed by Baur and
Adams in 1891. A major expedition in 1897, under the leadership of
C.M. Harris and including Rollo Beck, collected 59 Duncan tor
toises, of which 29 were delivered to Walter Rothschild, exhibited
at the London Zoological Conference in 1898, and later dispersed
among European zoos. Captain Noyes collected four Duncan tor
toises in 1900, and Beck collected five in 1901. The Academy expe
dition in 1905 and 1906 collected 86 tortoises from Duncan____25
males and 61 females. However, some time in the last two decades
of the nineteenth century, black rats escaped from a visiting ship
and quickly overran the island. Thereafter the rats ate every
hatchling tortoise as soon as it appeared above ground, and
natural replacement of the population ceased. One or two Duncan
tortoises were collected by many of the expeditions to the
Galapagos in the present century, but soon it became clear that
young animals were no longer present. William Beebe found only
one specimen in 1923, and he thought that he might have found the
very last individual. However, Townsend found tortoises still there
in 1928, and in fact one female Duncan tortoise collected by Town
send in that year survived in the Bronx Zoo, New York, until 1971,
at which point, old but still healthy, it was shipped back to the
Galapagos as a conservation gesture.
At the present time probably between 150 and 200 adult tor
toises still exist on Duncan Island. They have been exterminated in
the main caldera and in the central part of the island, but still per
sist on the southwestern slopes, where they are protected from
easy access from the sea by high cliffs. These tortoises, having had
no natural replacement this century, are very old and have smooth
shells with no signs of recent growth. Nevertheless, the females
are still laying fertile eggs, and since 1965 efforts have been made,
originally by Miguel Castro, former conservation officer of the
Charles Darwin Research Station, and subsequently by the Ecua
dorian National Park Service, to collect the eggs and complete the
370
incubation under artificial conditions. The tortoises are then main

tained for 4-5 years in captivity and taken back to Duncan Island
when they have attained a size at which they are no longer liable to
be eaten by rats. This program has been very successful, seventy-
one young tortoises now having been released on Duncan, some of
them in the central area from which tortoises had been extermina
ted long ago. The first twenty tortoises to be released were all
found to be doing well a year later, and all had shown good growth.
As an example of the growth rates of young Duncan tortoises
under natural conditions, three individuals which had respective
curved carapace lengths of 12.5”, 12.7”, and 11.8” when released
on Duncan on December 6, 1970, had reached lengths of 17.0”,
16.1”, and 16.6” when I found them on April 25, 1972.
Adult male Duncan tortoises are about 24” to 28” in straight
carapace length and have saddlebacked shells which tend to reach
their highest point either on the anterior marginals or on the first
central scute. The necks and limbs are long, and there are usually
whitish-yellow markings on the face and throat. The shell is nar
rowed anteriorly, but less so than in the Hood Island tortoises. The
anterior flanks of the shell, behind the reverted marginals, often
become encrusted with lichens. Females are smaller than the
males, usually being less than 24’ ’ long and weighing less than fifty
pounds. They are very flat and show little saddling of the cara
pace. All the mature females, and a few males also, show exten
sive pitting in the shell, where the bone has dissolved out and the
overlying scute area has subsided slightly into the gaps in the
bone. This is especially noticeable in the areas adjacent to the car
apace seams. Its more frequent appearance in females may be
associated with extraction of calcium from the shell for egg-shell
formation. , .
The Ecuadorian National Park Service officials have now
determined that the Duncan tortoise lays from 2 to 8 eggs, the
mean clutch size being 4.8. Fertility is high, around 80%. Nesting
takes place from August to December, and the natural incubation
period for six nests ranged from 85 to 120 days. Nests are allowed
to remain in situ until near to hatching time before being transpor
ted to the incubators at the Darwin Station, since premature dis
turbance of the eggs usually kills the developing embryo. The
hatchlings weigh about three ounces; while some are uniformly
black, others have large pale circles on each of the carapace
scutes. They show a strong tendency to flee from bright light, a
trait which would stand them in good stead under natural condi
tions, where exposed baby tortoises fall prey to hawks or get over
heated in the sun. The adults, too, hide under rock cairns through
out the heat of the day, venturing out in the cool of the morning and
evening to feed.
371
www.Ebook777.com
Red-checked mud turtle, Kinosternon scorpioides cruentatum, from Costa
Rica.
Head of the scorpion mud turtle, Kinosternon scorpioides. Photo by Russell

Mittermeier.
Carapace of the Mexican mud turtle, Kinosternon hirtipes.
The Big Bend mud turtle, Kinosternon hirtipes murrayi. Photo by Dr. R. W.
Barbour.
373
www.Ebook777.com
The Duncan Island tortoise is usually called Geochelone ele-
phantopus ephippium. However, the type specimen of Testudo
ephippium is a preserved male of uncertain origin in the Edin
burgh Museum of Science and Arts. This specimen is a saddleback
tortoise, 33” in straight length, which according to Gunther is a
relatively young specimen, still growing at the time of death. Gun
ther in 1896 compared the specimen with three Abingdon tortoises
and four from Duncan, and decided that agreement with the Dun
can specimens was perfect, while marked differences existed be
tween it and the Abingdon tortoises. However, subsequent findings
indicate that Gunther may well have been wrong. Of the hundreds
of Duncan tortoises collected since 1896, all were less than thirty
inches long, while the type of ephippium was still growing at 33”.
On the other hand, the Abingdon tortoise is known to reach a length
of at least 38”. Moreover, although the type of ephippium is
unlabeled it is known that Captain Basil Hall, who visited Abing
don but no other island in his Galapagos expedition in 1822, deposi
ted an Abingdon tortoise in the Edinburgh Museum, and this may
well be the same specimen. Also, the three abingdoni available to
Gunther were all, for some reason, exceedingly elderly specimens
with atypically thin carapaces. The specimens collected subse

quently show close similarity with the type of ephippium, and
indeed the agreement between Van Denburgh’s 32” male abing
doni (illustrated in Plate 27, Proc. Calif. Acad. Sci. 4th. Series Vol.
II Pt. 1) and the 33” type of ephippium (illustrated in Gunther’s
‘Gigantic Land Tortoises’, PI. XXXIX) is remarkably close. I feel
then that, strictly speaking, the name ephippium should refer to
the Abingdon tortoise and that a new name should be proposed for
the Duncan form.
The tortoises of Hood Island were collected on a massive scale
by the nineteenth century whalers. This island was the first one
that a ship approaching the archipelago from the south would
encounter; also the anchorage at Gardner Bay was quite secure,
and the tortoises were abundant, lived relatively close to the sea,
and were of a convenient small size for carrying. Townsend’s
analysis of whalers’ logbooks revealed that two ships visiting Hood
in 1831 carried off 585 tortoises between them, and many hundreds
374
more were removed in the following decade. However, the last big
take (67 tortoises) was made in 1847; at that point the population
apparently collapsed, and no search party thereafter was able to
locate more than seven tortoises. Extended searches by the
Academy party in 1906 revealed only three live animals. A 22.2”
young male from this collection was kept alive for a while in Gol
den Gate Park, San Francisco; it is now preserved in the Academy
of Sciences Collection, where it serves as the type specimen of
Geochelone elephantopus hoodensis.
Two more Hood tortoises were collected by the Pinchot expedi
tion in 1929, but apart from these almost none were found until the
inauguration of the Charles Darwin Research Station. Personnel
from the Station and from the National Park Service have now
located fourteen Hood tortoises after frequent searches of the
entire island—12 females and two males. The reasons for the stag
gered sex ratio are unknown, but fortunately the males in hand are
fertile and the fourteen individuals now constitute a captive breed
ing colony at the Darwin Station. After several years of experi
mentation, fertile eggs were at last produced in the 1970-71 breed
ing season, and at the time of writing the Station is raising about 30
healthy baby Hood Island tortoises. Craig MacFarland informs me
that the nests were made during the period July to December and
that the earlier clutches took much longer to hatch than the later
ones—probably because the weather is cooler during the later
months of the year than from January onward. The first clutch
took more than six months to hatch, while the last took less than
four months. The average clutch size of the Hood tortoise based on
21 nests at the Darwin Station is exactly six eggs.
The Hood tortoise is one of the smallest of the Galapagos tor
toises. No museum specimens of adult males are available for
measuring, but the larger adult male at the Darwin Station has a
carapace length of about thirty inches, and another adult male,
almost certainly from Hood Island, in the San Diego Zoo is of about
the same size. Mature females are only about 21-22 inches long.
The males have strikingly saddlebacked shells, strongly elevated
and narrowed behind the neck. The head is small, with extensive
white areas on the chin and throat, and the neck and limbs are
very long and thin. Females show the same characteristics, but in
a less extreme condition.
Eventually it is hoped that it will be possible to re-establish a
colony of tortoises on Hood Island. For such a move to succeed,
however, it would be necessary to complete the extermination of
the feral goats, which have overrun the island for many years and
which have denuded the vegetation to an extent that there is vir
tually no food for young tortoises. Once this has been done, it will
probably be desirable, at least for the initial stages, to enclose the
375
www.Ebook777.com
Head of the mud turtle Kinosternon acutum.
Plastral view of Herrera’s mud turtle, Kinosternon herrerai, a little-known,

presumably rare, species from eastern Mexico.
www.Ebook777.com
tortoises in a corral so that the population density will not be so low
that the tortoises will be unable to find each other. At the present
time, the population is so dispersed that copulation no longer takes
place, and all the females that have been found in recent years had
lichens on the upper rear parts of their shells—such growths are
rubbed off during copulation.
Of the three northern islands, Abingdon, Tower, and Bindloe,
only Abingdon, the northernmost and the highest, although not the
largest, has ever had a population of tortoises. The island is about
forty square miles in area, but much of this is fresh lava without
any vegetation and is consequently uninhabitable by tortoises.
Nevertheless, the relative accessibility of the vegetated parts of
the island had the result that the crews of whaling vessels in the
early nineteenth century were particularly ruthless in their collec
tion of tortoises from this island for provisioning their lengthy voy
ages to the Antarctic. Townsend records that a single ship, the
“Abigail,” collected no fewer than 142 tortoises from Abingdon on
one trip in 1837, and 67 were collected by the “Hector” in 1843.
Thirty-three were known to have been collected by two ships in
1848, but in that decade the population apparently reached a point
of collapse, and no subsequent ship’s crew was able to collect more
than six. Nevertheless, the absence of any introduced mammals
on Abingdon allowed a small population to continue to survive, and
in 1875 Commander Cookson collected four specimens, of which
three eventually found their way to the British Museum, where
they were designated as the types of Testitdo abingdoni. All three
were adult males, ranging from 34” to 38.5” in length and from 135
Outline of shell of Geochelone e. ab

ingdoni.
to 201 lbs. in weight. Very few more have been collected in sub
sequent years, but Rollo Beck collected a male and a female in
1901, which were deposited in the Tring Museum, and the California
Academy of Sciences party in 1906 collected three adult males and
an old shell of a fourth, ranging in length from 29.3 to 36 inches.
Townsend (1925) figures a live Abingdon tortoise which was exhib
ited at the Bronx Zoo, but gives no details about date or circum
stances of capture. Whatever population survived on the island in
the early part of the 20th century remained more or less undis
turbed from human interference until 1959, when an Ecuadorian
378
settler illegally introduced three goats to the island. These rapidly

multiplied to thousands and made serious inroads into the vegeta
tion. An expedition to Abingdon by Miguel Castro and David
Cavagnaro in 1964 spent nine days searching for signs of living tor
toises without success, although the skeletal remains of 28 individ
uals were found in holes and fissures. The conclusion made was
that the goats had so denuded the vegetation in the lower parts of
the island that the remaining tortoises had been forced to migrate
across unfamiliar ground in search of food and had all fallen into
crevices that lay, concealed by vegetation, across their path.
However, in 1971 a party of National Park wardens engaged in
goat extermination found some signs of living tortoises, and later
Joseph Vagvolgyi, a scientist based at the Darwin Station, found a
living tortoise which he reported to be of relatively small size. Not
realizing the significance of his find, he photographed the animal
but did not bring it back to the Darwin Station. In March 1972,
another goat-killing party actually located a young adult male
which was successfully translocated to an enclosure at the Darwin
Station. It was assumed until recently that this was a different
animal from the one found by Vagvolgyi, but examination of the
tortoise in hand and Vagvolgyi's photographs show them to be
identical. The possibility of finding a mate for the male thus
appears bleak, but the search continues. In 1972 the present author
found a complete carapace of an adult tortoise that had been killed
by man some years before, but was unable to find any signs of
living tortoises.
The type series of the Abingdon tortoise in the British Museum
indicates an extreme saddle-backed form with a very poorly ossi
fied carapace and a long neck with yellow areas on the jaws and
throat. The poor carapace ossification was used by Boulenger to
distinguish this race from the Duncan Island form, but the speci
mens subsequently collected show that the carapace, although
always saddle-backed, may be relatively thick. However, the Dun
can race is smaller (largest of 86 specimens collected in 1906 was
29.5” long), is less narrowed toward the front of the shell, has less
flaring carapace margins, and (usually) weaker scalloping of the
rear marginal scutes. Also, the Abingdon tortoise tends to reach its
highest point in the middle of the second vertebral scute, while the
Duncan race is usually highest at the very front of the shell. But
the character of the very short superior border of the eighth mar
ginal scutes of abingdoni, cited by Van Denburgh, is not constant.
Abingdon Island is unusual in having the combination of sad-
dlebacked tortoises and sufficient altitude to allow the growth of
moist, luxuriant vegetation in the interior. It appears, however,
that the tortoises were normally found not in the moist highlands,
but in the upper part of the arid zone, where competition for food
379
www.Ebook777.com
The common musk turtle or stink-pot, Sternotherus odoratus.
Plastral view of a razor-backed musk turtle, Sternotherus carinatus, showing

the extensive areas of soft skin between the plastral scutes.
380
Striped-necked musk turtle, Sternotherus minor peltifer.
Southern musk turtle, Sternotherus minor minor.
www.Ebook777.com
prompted evolution of the saddleback carapace shape as
described earlier. Despite the greater abundance of food and
water in the highlands, the necessity of crossing dangerous crevi
ces to reach suitable lowland breeding areas apparently prevented
colonization of the upper part of the island. The stomachs of those
tortoises collected on Abingdon in 1906 contained grass and cactus.
The Opuntia cacti on Abingdon still hold their succulent pads well
clear of the ground on eight-foot-high inedible trunks, out of reach
of the tortoises that have now all but disappeared, while on the
neighboring, tortoise-free islands of Bindloe and Tower, related
species of Opuntia assume the more familiar prostrate, spread
ing form.
The reproductive habits of the Abingdon tortoise are unknown.
Only one female tortoise has ever been collected by a scientific
party.
Tortoises were abundant on Chatham Island at least until the
mid-nineteenth century; whalers’ logbooks analyzed by Charles
Townsend showed that, of 10,373 tortoises recorded as collected in
the islands from 1831 to 1868, 4,326 were from Chatham. Charles
Darwin visited the island in 1835 and described the “Cyclopean
scene” of giant tortoises eating cactus, probably near the middle
of the island, judging by his description of the terrain. In the 1850’s
it was still possible for a whaling crew to collect over 300 tortoises
on a single visit, and Townsend reports a recorded total of 435 tor
toises collected on Chatham in 1861. Nevertheless, in the following
decades the depredations of the whalers, settlers, and introduced
mammals of various kinds made serious inroads into the tortoise
population. In 1906, The Academy party was only able to find one
tortoise on Chatham—an ancient but small female with one foreleg
missing. Several shells and skeletons were also collected, one of
which became the type specimen of Testudo chathamensis (now
known as Geochelone elephantopus chathamensis), described by
Van Denburgh in 1907. In his 1914 paper on Galapagos tortoises,
Van Denburgh assumed that the Chatham tortoise was almost or
quite extinct, and this belief prevailed for many years. However,
although the tortoises have by now almost certainly disappeared
from the entire southwestern half of Chatham, evidence began to
accumulate in the 1960’s that a few tortoises still survived in the
uninhabited, arid, and rarely-visited northeastern end of the
island, which is largely separated from the fertile inhabited parts
by massive fields of fresh lava. In 1965 Miguel Castro and two com
panions actually managed to locate four tortoises and concluded
that there might be as many as sixty still left on the island. Eight
were found on a second expedition in 1967, and finally, in 1969, Cas
tro, Juan Black, and Oswaldo Chappi found 57, bringing the grand
total of marked animals to 69. It is now estimated that between 450
382
and 500 tortoises exist in a rather restricted area between Pan de

Azucar and Cerro Media Luna in the northeastern part of Chat
ham ; 213 of these have now been marked. The area is very hard to
locate—almost impossible without a guide, since the terrain is
very broken and trackless. However, I was privileged to be taken
to the tortoise area in April 1972 by wardens from the Park Ser
vice, and in the course of a single morning our party located 91 tor
toises. We also noticed that the sex ratio of the tortoises was reas
onably close to equality. Nevertheless the future of this population
is far from assured; the population is not as senile as that of, say,
Duncan Island, but nevertheless very few young specimens exist,
and none can be found of sizes corresponding to ages of less than
about fifteen years. The reasons for the animals’ failure to repro
duce in recent years are manifold; it is probable that the numer
ous feral donkeys in the area destroy the buried eggs with their
hooves, and those young which do hatch are eaten by feral dogs.
The habitat is also occupied by feral cats and small numbers of
goats and cattle, which may have additional adverse effects.
The Chatham tortoise is a small to medium-sized race; the
biggest males reach a carapace length of about 36”, but females do
not greatly exceed 24”. Young specimens have relatively low
shells which are not narrowed in front. The highest point of the
Outline of shell of Geochelone e.

chathamensis.
shell in adults is usually in the middle of the second vertebral

scute. The margin of the shell above the hind limbs is strongly
flared in both sexes, and the anterior marginals are frequently
reverted. Fully adult males have the “saddleback” shell shape,
narrowed and elevated in front, with the front pair of marginals
forming a straight line in anterior aspect, sloping sharply down
ward at the junction with the second marginals. The widest point
of the shell in old males is situated across the posterior flaring of
the carapace, but in younger males and in females the widest point
is near the middle. Van Denburgh noticed from the very few indi
viduals available to him that the pectoral scutes of chathamensis
are separated on the midline. A larger series of eighteen live speci
mens that I examined in 1972 showed pectoral separation in exact
ly 50% of cases, these nine specimens including all situations from
complete failure of either pectoral to meet the midline to cases in
which one pectoral met the midline and the other nearly did. But
383
www.Ebook777.com
Flat musk turtle, Sternotherus minor depressus, a rare and localized sub
species from northern Alabama.
Plastral view of a hatchling giant musk turtle, Staurotypus triporcatus, from

Veracruz, Mexico.
384
Mexican giant musk turtle, Staurotypus salvini, from the Pacific drainage of
southern Mexico, Guatemala, and El Salvador.
Narrow-bridged mud turtle, Claudius angustatus, from tropical eastern Mex

ico.
385
www.Ebook777.com
despite the inconstancy of this condition, it is not found, except
possibly as a very rare variant, in other Galapagos tortoises, ex
cept possibly those of Abingdon and Duncan.
In color the Chatham tortoises are the usual uniform black,
but generally there are white or yellowish markings around the
chin and comers of the mouth and frequently around the nostrils.
The Chatham tortoises are reported to begin nesting in September,
and typical nests number from four to six eggs. They show the
typical giant tortoise propensity to concentrate in muddy pools
during the rainy season and to eat virtually any green vegetation
they may encounter.
Only a single tortoise has ever been collected on Narborough
Island. This was an adult male, found by Rollo Beck on April 3,
1906. This animal, the type specimen of Geochelone elephantopus
phantastica, is now in the California Academy of Sciences. It
measures 34.5 inches in straight length and was found about half
way up the volcano on Narborough Island. The tortoise shows an
extreme saddlebacked condition, with very strongly reverted an
terior marginals, and yellow markings on the head and throat.
Beck also found a few old tortoise droppings on Narborough, which
showed that the animal had been feeding upon the very spinose
Cereus (now known as Jasminocereus) cactus.
Since Beck’s discovery of the type specimen, the only evidence
of living tortoises on Narborough Island was that found by John
Hendrickson in 1964, which consisted of some old droppings and
Opuntia cactus pads with bite-shaped sections missing. In subse
quent years several parties have made thorough searches of the
southern slopes of Narborough, finding many land iguanas and
what looked like excellent tortoise habitat, but without finding any
sign of living tortoises. Nevertheless, the area is large and the ter
rain between the “islands” of vegetation exceedingly difficult, so
it seems not unlikely that a few individuals may still survive some
where on the island. But the population density is so low that even
in 1906, Beck’s specimen was apparently unable to locate females
and had resorted to attempted copulations with a rock on the trail.
It is surprising that the Narborough tortoise should be so ex
cessively rare, since the island has never been inhabited by man
and remains to this day free of feral mammals. Moreover, none of
the whalers’ logbooks ever mentioned finding tortoises on this
island. One can only assume that the recurrent vulcanism of the
island has either killed most of the tortoises or has separated the
breeding areas from the feeding areas so that only a relictual, non
breeding group remains. It is unlikely that Beck’s specimen was
an introduction, since not only was it found a long way from the
coast, but in shape it does not check out with any race from any
other island.
386

phantastica.
Barrington Island is a rather small island with no humid zone

which at the present time is not inhabited by tortoises. However,
Townsend found one whaleship record of 22 tortoises having been
collected on this island in 1839. One Captain Levick informed the
Academy party in 1906 that thirty years earlier tortoises were
found scattered all over Barrington, but that he had removed them
all. However, this informant was probably unreliable, since much
of the other information he had given about Galapagos tortoises
was definitely untrue. The Academy party found none alive them
selves, but did find fragmentary remains of about fourteen indi
viduals and two old semi-fossil eggs.
Approximately six tortoises, probably from Indefatigable
Island, were deposited on Barrington in 1956, and at least one of
these is still there.
Tortoises were formerly abundant on Charles Island, but they
are now definitely extinct. Captain Porter collected 400-500 tor
toises on Charles in 1812, and Townsend found logbook records of
234 being removed in 1831, at least 226 in 1832, 570 in 1834, 90 in 1835,
and 24 in 1837. The island had been inhabited by political prisoners
exiled from Ecuador for six years when Charles Darwin visited the
island in 1835, and Darwin wrote of the tortoises that:
.. their numbers have of course been greatly reduced
on this island, but the people yet count on two days’ hunting
giving them food for the rest of the week. ”
Nevertheless we have no evidence that live tortoises could be
found on Charles Island later than 1837, and B. Seemann, the
naturalist on board the “Herald,” which visited Charles Island in
1846, stated that there were no longer any tortoises living on that
island.
Very few specimens of the Charles Island tortoise collected
during the lifetime of the race are in existence; but there is a male
in the collection of the Boston Society of Natural History which was
collected by Commander John Downes, of the U.S. Frigate “Poto
mac” in 1833. There is also a well-mounted specimen on exhibition
in the British Museum (Natural History). Moreover, the American
Museum of Natural History has a good series of skeletons of this
387
www.Ebook777.com
Two views of the Zambezi soft-shelled turtle, Cycloderma frenatum, from
Lake Malawi, Malawi. Photos by Dr. W. E. Burgess.
Trionyx ferox, the Florida soft-shell, is the largest New World trionychid.
389
www.Ebook777.com
species, the result of a fortunate discovery by Townsend in the
mid-1920’s of a cave in an arid part of the island into which tor
toises had fallen and eventually died. The discovery of this cave is
described in a paper by Robert Broom (Zoologica 1929: IX; 8: 306-
309) as follows:
“It would be difficult to imagine a more effective trap
for tortoises than the well-like entrance to the cave on
Charles Island from which the skeletons were taken.
“It must have operated automatically as a death trap
for centuries. The brushy half-concealed entrance is merely
a hole in the ground a dozen feet in diameter and 20 feet
deep. With a steep slope at one side, the unlucky tortoise that
tumbled in did not necessarily strike bottom with a fatal
crash, but rather rolled down an incline it could not ascend.
“The rocky floor of the cave is not wide but leads into a
few low passages under the lava, all strewn with dry bones
of tortoises that had crept everywhere in search of an outlet.
The brittle remains of the earlier victims had been crawled
over repeatedly and gradually broken up by those that were
trapped subsequently from time to time.
“It is not likely that the latest and best-preserved of
these entered the cave less than 90 years ago. According to
the logbook records of 79 whaleships examined by the
writer, the last tortoises were taken from Charles Island in
1837.
“The bleached and bony remains of those not too anti
quated and fragile to be removed, had long lost their dark
homy plates, which lay curled and twisted beside them. In a
dozen of these, both carapace and plastron were practically
intact, while skulls and leg bones had usually been disturbed
and scattered. A considerable amount of broken tortoise
remains had long since become mixed with the soil of the
cave floor. The later arrivals lay where they had died, their
large white carapaces showing conspicuously as our flash
lights were turned in their direction. ”
It was formerly considered that the original type specimen of
the Galapagos tortoise, Richard Harlan’s specimen which he
named Testudo elephantopus, was a Charles Island tortoise. How
ever, it now appears that Harlan’s specimen (of which only frag
ments still exist) was more probably from Cerro Azul on Albe
marle Island, so that, under modem nomenclature, the Charles
tortoise becomes Geochelone elephantopus galapagoensis (Baur
1889), while the Cerro Azul race becomes Geochelone elephanto
pus elephantopus (Harlan 1827). The Charles Island tortoise, from
the available specimens, appears to have been a race of small to
390
Specimen of Geochelone e. galapagoensis in the British Museum (Natural

History).
medium size, with an elevated, “saddleback” anterior carapace

margin, but lacking the extreme anterior narrowing of some of the
saddleback races.
There is no satisfactory evidence that tortoises ever existed
naturally on Jervis Island, a tiny island only about two miles in
diameter. The only whaleship which is recorded as having visited
this island looking for tortoises was the “Zena’s Coffin,” which
searched Jervis on July 12, 1850 without finding a single tortoise.
Moreover, at this time Jervis Island had no feral mammals and
had not been despoiled by man, so that one would have expected
that if tortoises were there at all, they would be abundant. The
Academy expedition did find a single live tortoise on Jervis on
December 20, 1905, and Van Denburgh found that this specimen
compared very closely with the type specimen of Testudo walla-
cei, a shell of unknown origin in the Rothschild Museum at Tring.
However, Van Denburgh also quotes a “rumor” that Georg Baur
had introduced tortoises to Jervis Island from southern Albe
marle, and since the specimen collected on Jervis in 1905 agreed
almost exactly with the characters of the southern Albemarle tor
toises, I am inclined to accept this rumor as true. It is on definite
391
www.Ebook777.com
A Gulf Coast soft-shelled turtle, Trionyx spiniferasper, from Atlanta, Georgia.
Texas soft-shelled turtle, Trionyx spinifer emoryi, a very abundant turtle in

aquatic situations in southern Texas.
392
A juvenile smooth soft-shelled turtle, Trionyx muticus calvatus, from the

Conecuh River of southwestern Alabama.
The peacock soft-shelled turtle, Trionyx hurum, a rare and beautiful species
from the Indus and Ganges Rivers. Photo by Walter Sachsse.
www.Ebook777.com
wallacei.
record that Baur found tortoises on southern Albemarle when he

visited the archipelago in 1891. A party collecting tortoises for
Rothschild in 1897 found the track of a single tortoise near the
lagoon on Jervis, and this may have been a specimen brought over
by Baur and later collected by the Academy party. Its presence
near the landing place rather than in the interior of the island
would lend support to this hypothesis.
GENUS TESTUDO
At one time all turtles—land, freshwater, and marine—were
included in the genus Testudo. However, in 1805 Brongniart pro
posed the new genera Chelonia (for the marine forms) and Emys
(for the freshwater forms), and Bell in 1828 narrowed the use of the
name Testudo further by the separation of those tortoises with
hinged shells, which he placed in the new genera Kinixys and
Pyxis. Testudo thereafter remained a “catch-all” genus for all tor
toises without some sort of obvious peculiarity until quite recently,
when Ernest Williams (in Loveridge and Williams, 1957: Bull.
Mus. Comp. Zool. Vol. 115, no. 6) revived the long-neglected genus
Geochelone Fitzinger 1825, which he divided into several sub
genera, and restricted Testudo to the five obviously closely related
circum-Mediterranean tortoise species. Testudo under this new
restriction was defined as follows:
“Skull with triturating surface of maxilla moderately or
weakly ridged or without ridging; median premaxillary
ridge absent; maxillary not entering roof of palate; anterior
palatine foramina small, concealed or large and conspic
uous; pro-otic typically concealed dorsally and anteriorly by
parietal; quadrate enclosing stapes or not; surangular sub
equal in height to prearticular; neck with third or fourth
centrum biconvex. 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 fourth; submarginal scute absent; frequently a
single suprapygal, if two, then typically separated by a
straight transverse suture. Plastron with posterior lobe
more or less hinged in one or both sexes; gular region but
394
little thickened or produced; gulars paired, longer than

broad. ”
Williams’ new definition of the genus Testudo has gained gen
eral acceptance, and we shall use it here.
Testudo graeca, commonly known as the Moorish tortoise or
the Mediterranean spur-thighed tortoise, is found in North Africa,
the Near East, and extreme southeastern and southwestern
Europe. The species may be recognized by the moderately domed
carapace, of a dull yellow or olive color, with black anterior and
lateral margins to each of the scutes and also small irregular
black markings near the areola of each of the carapace scutes. The
precise extent of the black pigmentation is very variable. The
nuchal scute is narrow but slightly wider posteriorly than anterior
ly. The supracaudal scute is undivided. The plastron is yellow or
greenish-yellow, with a large black blotch on each of the larger
scutes. In mature females the posterior lobe of the plastron is
slightly moveable. The head is of medium size and is dorsally
covered with numerous scales of which the only large one is the
frontal, which too is sometimes fragmented. A single prefrontal is
present, and paired supranasals. The jaw surfaces are not serrate,
but the beak is weakly hooked and is tricuspid. The forelimbs each
bear five claws, and their dorsal surfaces are covered with
numerous heavy, pointed scales which form 3 to 6 longitudinal
rows and 4 to 7 transverse rows (from the elbow to the outer claw).
There is a prominent spur in the thigh region on each side.
Large adult Testudo

graeca found by the
author crossing a high
way near Istanbul, Turkey.
www.Ebook777.com
Trionyx subplanus, a small soft-shell from Thailand.
Black-rayed soft-shelled turtle, Trionyx cartilagineus, from Southeast Asia.
396
The Chinese soft-shelled turtle, Trionyx sinensis, is widely used for food in
the Orient and has been introduced into Hawaii.
the narrow-headed soft-shell, a gigantic species from northern

Chitra indica,
India, Thailand, and Malaysia. Photo by Dr. Sherman Minton.
397
www.Ebook777.com
Four subspecies of the spur-thighed tortoise are currently con
sidered valid: Testudo graeca graeca in extreme southern Spain,
the Pityusen Islands, much of Morocco, and the coastal parts of
Algeria, Tunisia, and Libya as far east as Cyrenaika; Testudo
graeca ibera in southeastern Europe (Turkey, Bulgaria, and
northeastern Greece), through Asia Minor to central Iran; Tes
tudo graeca zarudnyi in eastern Iran; and Testudo graeca terres-
tris in Syria, Israel, Sinai, northern Egypt, and northeastern
Libya. The African races, graeca and terrestris, are smaller than
ibera and zarudnyi, rarely exceeding eight inches in carapace
length, while ibera and zarudnyi often reach shell lengths of about
eleven inches. The head of graeca is dark and the carapace is rela
tively depressed and elongate; terrestris has yellow on the sides
and top of the head, and the carapace is strongly elevated. The
Asiatic races ibera and zarudnyi typically have relatively indis
tinct markings, especially zarudnyi in which the ground color is so
dark that markings are barely apparent. This race is also distin
guished by the more elongate carapace, of which the hind margin
is slightly serrated. It is worthy of note, incidentally, that Testudo
graeca does not receive its specific name from any reference to
Greece, but rather from the appearance of the scutes of the cara
pace, which struck Linnaeus as looking like a Greek mosaic. More
over, T.g. ibera is not named after the Iberian Peninsula, but
rather after Iberia in Transcaucasia.
It is the spur-thighed tortoise that supplies the bulk of the pet
shop tortoises in Britain. The tortoises are for the most part of
Moroccan origin, and M.R.K. Lambert, who recently conducted a
study of this trade for the World Wildlife Fund, estimated that
more than 300,000 tortoises are shipped from Morocco to Britain
each year. Lambert found that tortoises had become scarce in
many areas where they had formerly been abundant, but even in
those areas where they were now scarce collecting still continued,
since the collectors were primarily shepherds for whom tortoise
collecting was merely a sideline to their normal work. Lambert
estimated that about 5 million tortoises remained in the entire
country—not a large number considering the size of the country
and the number exported each year. Only tortoises with plastral
lengths of between 4 and 6 inches are exported, tortoises smaller
than this being legally protected and larger than this undesirable
to the trade because of heavy shipping costs and also because of
the preference shown by the ultimate buyers for smaller tortoises.
However, tortoises less than four inches long, or more than six
inches, or those of intermediate size but whose shells are damaged
and thus not suitable for the pet trade, are killed in some numbers
for use of their shells as sounding boxes for crude, locally made
“banjos.” But the banjo trade is small compared to the trade in
398
Testudo graeca terrestris occurs in the Near East and northeastern Africa.
live animals. The immediate result of heavy collecting of “legal-

size” animals is to leave almost entirely very small males and
particularly large females, a combination that may well result in
decreased mating success.
Loveridge and Williams described two spur-thighed tortoises
of Algerian origin that hatched in September 1905, at which time
they measured 1.85” and 1.93” over the curve. Growth was steady
for the next 39 years as the turtles were raised by Mr. C.F. Moysey
in an English garden. In 1944 the tortoises had over-curve meas
urements of 11” and 14-3/8”, the straight length of the latter speci
men being 10-7/8”. By 1955 the larger tortoise had the same cara
pace length, and a weight of 4.252 kg—thought by Loveridge and
Williams to be a record for North African graeca.
Testudo graeca hibernates during the cold months but
emerges as early as February in the hot coastal areas, and indi
vidual tortoises may emerge temporarily during warm days even
in the depth of winter. Courtship takes place in the spring and in
volves immobilization of the female by repeated butting, as well as
biting at the head and forelimbs. During copulation the male gapes
widely and emits a muffled whistling cry. Eggs are normally laid
399
www.Ebook777.com
Juvenile Fly River turtle, Carettochelys insculpta, photographed at the San
Antonio Zoo.
Adult female Fly River turtle, Carettochelys insculpta.
400
Central American river

turtle, Dermatemys mawi,
from the Caribbean low
lands of southern Mexico
and Guatemala.
Chinese big-headed turtle, Platysternon magacephalum megacephalum.
401
www.Ebook777.com
in May and June and hatch in September or October. In North
Africa, it appears that the average clutch numbers only two or
three eggs. The size of the egg is variable, probably depending at
least partially on the size of the female; a small egg measures
1 x 1.2”, a large one 1.2” x 1.4”. Sometimes eggs are more
strongly ellipsoidal than these figures would suggest.
Testudo graeca is essentially vegetarian and in captivity
shows a great fondness for lettuce and fleshy-leaved plants, as well
as yellow flowers. There is a lot of idiosyncrasy among captive
specimens in the foods they will accept; for example, some are
very fond of milk-soaked bread while others will not touch it.
The majority of the spur-thighed tortoises imported into
Britain die within the year, the climate being too cold and damp.
However, specimens that are allowed the run of large gardens in
southern England often survive for many years. A tortoise that
originally belonged to Archbishop Laud survived from 1633 to at
least 1730, and possibly 1753; its remains are still preserved at
Lambeth Palace. Another is supposed to have lived 96 years in a
garden in Cornwall.
The habits of Testudo graeca terrestris do not appear to differ
markedly from those of the nominate race. Eggs in Israel are laid
in June or July and number 3 to 5 per nest. The eggs measure about
1.2” x 1.6”. The hatchling is about 1.2” in carapace length and is of
an overall dull yellow color except for a dark spot on each verte
bral scute. As with all races of this species, the habitat embraces
both lowland and mountainous country, but the population densi
ties are much higher at lower altitudes. James Buskirk found that
mountain tortoises in Israel had a much higher incidence of shell
damage than those from lowland areas.
The Near East subspecies T.g. ibera reaches consistently lar
ger adult sizes than graeca, adults often being over ten inches
long. The record for the race is 12”, for a specimen from Iglitza,
on the Danube. I have witnessed copulation of this race in the wild
on two occasions in central Iran, once on August 23, once on Sep
tember 10. The male showed essentially the same butting, mount
ing, and calling behavior described above for T. graeca graeca; in
addition, the female in one of the cases responded to the male’s
advances by walking away and shaking her head vigorously from
side to side. An egg that I found inside a dead tortoise in Iran
measured 1.4” x 1.6”.
The habits of the eastern subspecies T.g. zarudnyi have not
been described. I once found six specimens that I referred to this
subspecies, all hiding under rocky ledges beside a stream north
east of Isfahan, Iran.
Hermann’s tortoise, Testudo hermanni, is, most confusingly,
often referred to as Testudo graeca in the older literature. This is a
402
Plastral view of Testudo her-

manni robertmertensi from
southwestern Europe. Photo
by Dr. Robert Mertens.
strictly European species; the typical race, T. hermanni herman

ni, is found in southern Italy, Sicily, Yugoslavia, Albania, Greece,
the Aegean Islands, Bulgaria, European Turkey, and southwes
tern Rumania. It is apparently much more abundant at sea level
than at high altitudes. Testudo hermanni robertmertensi is found
in west-central and northwestern Italy, southern France, north
eastern Spain, Corsica, Sardinia, and the Balearic Islands.
Testudo hermanni differs from T. graeca in lacking the thigh
tubercles, having a divided supracaudal scute (though the division
maybe apparent only on the outside of the scute), and in reaching
a smaller adult size. A long, narrow nuchal scute is present. The
carapace is moderately domed, pale yellow in ground color, with
extensive black areas covering the anterior and lower part of each
marginal scute and the anterior and lateral parts of the costals and
vertebrals. The plastron is pale yellow with a black band of vari
able width along each side, which in many specimens is interrup
ted at the transverse seams. The plastron is not hinged. There is a
horny claw-like tip to the tail in both sexes. The sexes show the
usual differences; the tail of the male is longer and thicker than
that of the female, and the male’s plastron is distinctly concave
and posteriorly truncated. The subspecies are distinguished by the
more elevated carapace and more strongly contrasting shell pat
tern of robertmertensi and by the presence of a light spot behind
403
www.Ebook777.com
404
www.Ebook777.com
and beneath each eye. It is possible that this race is smaller than
T.h. hermanni; K. Heron (Int. Turt. Tort. Soc. J., 1968, No. 1 , p. 18
et seq.) examined over 400 French'specimens and concluded that a
large female reaches a length of a little over six inches, and a large
male about an inch less. The largest specimen encountered,
apparently an escaped pet, measured only 6ty”.
We are indebted to K. Heron for a number of excellent obser
vations on T. hermanni robertmertensi in a garden on the south
coast of France. The tortoises awoke from hibernation in March
and had two activity periods, one lasting until June and the other
until hibernation in November. Courtship took place all summer,
and eggs were generally laid in May and June, while hatching took
place in August and September. Some individual tortoises were
recorded to have laid twice within a season, usually at intervals of
between 19 and 23 days, but on one occasion the second clutch was
deposited after an interval of 31 days. Of forty nests, one (made by
the largest female) contained five eggs, six contained four eggs, 22
contained three eggs, six contained two eggs, and five contained
single eggs. The eggs are hard-shelled, slightly elongate, and have
a pinkish cast when newly laid. Incubation time ranges from 73 to
104 days. Rainy weather during incubation would slow the develop
ment, but rain near hatching time would facilitate the ascent of the
newly-hatched young to the surface, and during heavy rains hatch
lings might appear from several nests laid at different times.
A female ready to nest appears very active and energetic.
Once excavation of the egg cavity was commenced, it would be
completed unless the tortoise was disturbed or encountered an
immoveable root or stone. Excavation took from one hour to
hours and was never observed to be facilitated by the discharge of
urine to improve the clinging quality of the soil. Excavation is car
ried out by alternating movements of the two hind feet, and once a
hole of adequate depth has been constructed, it is enlarged anteri
orly into a rough flask-shape. The eggs are laid at about five-min
ute intervals, and each one is placed in position with the hind feet.
After a variable and often lengthy period of shoving and patting,
the former excavating action of the hind legs is reversed and the
eggs are buried.
One female tortoise was first observed nesting when its age
was known to be 7.75 years. The hatchlings are shades of grey
rather than black and yellow, and weigh about 10 grams.
Under captive conditions, Hermann’s tortoises often develop
abnormal shells, the scutes becoming freakishly humped and the
2nd to 4th central scutes appear to grow unnaturally slowly com
pared to the others. This species in captivity is highly omnivorous,
happily eating vegetables such as lettuce and radishes or almost
406
any kind of fruit, but also relishing canned dog food, ham rind,
cheese, and chicken bones.
The margined tortoise, Testudo marginata, is the true
“Greek” tortoise, being restricted to southern Greece, from the
Taygetos Mountains to Mount Olympus, although Walter Hellmich
(Reptiles and Amphibians of Europe) writes that “it is thought
that monks introduced them into Italy, and they have been discov
ered recently in Sardinia.” This is the largest member of the
genus, adults reaching a carapace length of a little over one foot.
Young specimens look very similar to T. graeca, although they
lack the thigh tubercles, but with age this tortoise develops a very
distinctive serrated flange around the hind margin of the cara
pace, caused by a great elongation and flaring of the hind margi
nal scutes. The overall coloration is very dark, the carapace of the
adult being dark brown or black except for light areolae on the
central and costal scutes and triangular or irregular light mark
ings on the marginals. The plastron is basically yellow, but each of
the larger scutes has a large triangular dark figure. The head and
limbs are heavily pigmented, and the upper jaw is strongly
hooked. There are numerous enlarged scales on the forelimbs. The
alveolar ridge of the upper jaw is rather short and feeble. The head
A pair of Testudo marginata at the Atlanta Zoo, Georgia. Notice the triangular
dark spots and the flaring posterior carapace margin.
407
www.Ebook777.com
Femsile green turtle, Chelonia mydas, returning to the sea. Photo by Keith
Gillett from The Australian Great Barrier Reef in Color.
408
Galapagos green turtle, Chelonia mydas agassizi, on the nesting beach.
Juvenile flatback sea turtle, Chelonia depressa, from northern Australia.
409
www.Ebook777.com
Hatching of the margined tortoise, Testudo marginata. Photos by Othmar
Stemmier.
scalation includes a large prefrontal followed by a frontal scale of

nearly equal size. The habits of the margined tortoise do not
appear to have been studied.
Testudo horsfieldi, commonly known as Horsfield’s tortoise or
the Afghan tortoise, is the most eastern representative of the
genus, being found from the northeast shores of the Caspian Sea
eastward through Kazakhstan to Lake Zaysan and thence south-
westward to Waziristan and eastern Iran. The species is wide
spread in Afghanistan, and in Pakistan is known from northern
and western Baluchistan. There is an isolated record for just south
of Kuybyshev, on the Volga. This is the only European record for
the species and, at 52° 30’N, is the northernmost record for any tor
toise. This is a fairly small species, normally considered adult at
six to eight inches in carapace length, although the type of Testudo
baluchiorum, now considered a synonym of T. horsfieldi, had an
abnormally domed carapace 8.65 inches in length. Moreover, two
hatchlings obtained by Dick Goergen of East Amherst, N.Y., in
1969 had by August 1,1973 reached shell lengths of 9-5/8” and 9%”
respectively, with weights of 4.06 lb. and 4.14 lb.
The Afghan tortoise has a broad, rounded, depressed cara
pace, distinctly flattened in the vertebral region. Some specimens
have a uniformly light tan carapace, but more commonly there is
410
extensive black pigmentation which may on occasion cover vir

tually the entire shell. It has not yet been demonstrated if these dif
ferences in pigmentation show geographic correlation. The nuchal
scute is long and narrow, and the vertebral scutes are consider
ably broader than long. The plastron is large, truncate or emargi-
nate anteriorly, and notched posteriorly. Each of the plastral
scutes is usually marked with a large black blotch. The head is
rather small, with symmetrical scales on the upper surface; the
upper jaw is tricuspid anteriorly, the lower jaw singly cusped. The
front limbs are strong, have relatively immoveable wrists, and
each is equipped with four well-developed spiky claws. All other
members of the genus have five claws on each forefoot.
The habits of the Afghan tortoise have not been thoroughly
studied. Sherman Minton found the species in Pakistan to inhabit
altitudes of 5000 to 7000 feet and to be especially abundant in grassy
places near springs or small streams in hilly, rocky terrain; at one
time he had seven specimens in view simultaneously in such an
area. In the hot months of May and June the reptiles took refuge in
shallow burrows beneath large boulders and outcrops of rock dur
ing the heat of the day, emerging to feed during the early morning
and evening. Minton found an egg of the Afghan tortoise to meas
ure 41 x 28 mm. On the other hand, Malcolm Smith reports that a
female killed toward the end of May contained five fully formed
eggs, 50 x 35 mm in size. Both of these writers mentioned that the
Afghan tortoise rarely eats grass, preferring flowers and fruits
and the thick fleshy leaves of various plants.
The rounded carapace of

the Afghan tortoise, Tes
tudo horsfieldi, is dis
tinctive. Photo by
Dr. Herbert R. Axelrod.
www.Ebook777.com
412
A loggerhead turtle, Caretta caretta, photographed by Keith Gillett, from The Australian Great Barrier Reef in Color.
Plastral view of a very young Kemp’s ridley turtle, Lepidochelys kempi.
Nesting female Kemp’s ridley turtle, Lepidochelys kempi, the most im
minently threatened of the seven sea turtle species.
www.Ebook777.com
Winters can be very cold in the arid steppe highlands that con
stitute much of the range of the Afghan tortoise, and it is interest
ing to speculate as to how the animals avoid freezing conditions.
There is no doubt that this species has sufficient “anti-freeze” in
its body fluids to survive a remarkable degree of cold; Mertens
records that it can tolerate body temperatures as low as 23.3° F.
Moreover, the depressed, gopher-like shell of the Afghan tortoise,
as well as its strong forelimbs and well-developed, relatively im
mobile claws, suggest that it may well dig extensive burrows and
thus escape exposure to extreme cold.
The little Egyptian tortoise, Testudo kleinmanni, although ap
parently related to and probably derived from Testudo graeca, is
considered sufficiently distinct to be placed in a separate sub
genus, Pseudotestudo. This subgenus is defined on the basis of the
unridged maxillary bones, the large, exposed anterior palatine
foramina, and the quadrate not enclosing the stapes. Other
features include the single supracaudal scute, the moveable hind
plastral lobe of both sexes, and the absence of a claw on the tip of
the tail.
The Egyptian tortoise, Testudo kleinmanni, has a moveable hind plastron

lobe.
414
Testudo kleinmanni looks

much like Testudo graeca
from above but has a wider
nuchal.
Testudo kleinmanni is found in western Cyrenaica, Libya, and

northern Egypt, including northern Sinai. The species is very
restricted in Egypt, only being known from the northwestern
deserts of the Nile delta and the region of the Suez Canal. This is a
very small species, males reaching a carapace length of about
four inches, females up to about five inches. The carapace is
highly domed and is almost uniformly pale yellowish or yellow-
green in color, often with narrow black edging to the scutes, but
without any dark pigmentation near the areolae. The plastron is
immaculate except for a single dark blotch on each of the abdomi
nal scutes. The nuchal scute is relatively large and much wider
posteriorly than anteriorly, and the carapace is distinctly emargi-
nate above the head. The limbs and head are usually immaculate
yellow, contrasting strongly with the jet black eyes; but some
times there is a dark spot on the crown of the head. The beak is tri
cuspid; the large frontal scale is often at least partially frag
mented. Many of the scales on the forelimbs are very large. The
supracaudal scute is occasionally divided and projects beyond the
marginals. There are no enlarged tubercles on the hind side of the
thigh, but each heel has a large conical spur-like tubercle.
Stanley Flower reports that the Egyptian tortoise, in captivity
in Cairo, mates in September and October. The mating male emits
sounds that have been likened to the intermittent winding-up of a
metal spring. Eggs, which measure about 1.2” x 0.9”, are reported
to be laid in May and June. The hatchlings, which are completely
yellow, emerge mostly in October and November.
415
www.Ebook777.com
Olive ridley turtle, Lepidochelys olivacea, nesting in Surinam.
Female Atlantic hawksbill turtle, Eretmochelys imbricata, climbing the

nesting beach in French Guiana.
416
Two views of the eastern Pacific hawksbill turtle, Eretmochelys imbricata,

from the Galapagos Islands.
www.Ebook777.com
In captivity this species requires high temperatures and low
humidity, hut it thrives when properly maintained, and there is
one record of a captive specimen surviving for 21 years and 25
days.
GENUS GOPHERUS
This is a purely North American genus, with four species. The
genus is defined on the basis of several characters, including the
modification of the forelimbs for digging (the forelimbs are flat
ter, from front to back, and are more rigid in the area of the
“wrists” and “hands” than those of other tortoises) and by the
presence of a median ridge on the premaxilla. A broad nuchal
scute is present; the carapace is usually flattened, and the gular
scutes are often extended forward into a fork or blunt projection.
The neural bones show a pattern of alternating octagons and
squares, and the lower ends of the pleural bones show a strong al
ternation in width.
The genus Gopherus is thought to have been derived from the
extinct North American genus Stylemys about 50 million years
ago, and in former times it was very widespread throughout the
warmer parts of the North American continent. Climatic changes
have now eliminated the larger species from the United States and
from all but a very small area in north-central Mexico, while the
three smaller surviving species too appear to be on the retreat and
now have relatively small and completely isolated ranges in the
southeastern, south-central, and southwestern United States.
The desert tortoise, Gopherus agassizi, is found in southeas
tern California, western Arizona, and extreme southern Nevada
and southeastern Utah. In Mexico the range extends for several
hundred miles along the coastal lowlands of Sonora, about as far
south as Alamos. It is a medium-sized species, large adults being
about eleven inches long, while the record specimen, a freakishly
large long-term captive individual, measured about seventeen
inches. The species can be distinguished by the relatively narrow
head, with the alveolar ridges of the upper jaw meeting at a sharp
angle, and the large hind foot. The latter feature presumably
assists locomotion on soft sand.
The desert tortoise has a uniformly brown carapace when
mature, rather low in profile, with more or less parallel (some
times slightly concave or convex) sides. The hind margin of the
shell is somewhat serrated and the carapace scutes usually show
well-defined growth annuli Young specimens have the central
part of each carapace scute lighter in color than the margin. As
with all members of the genus, there is a pair of “scent glands”
under the chin.
418
The desert tortoise is found over a wide range of latitudes and

altitudes, and although no subspecies have been described, the
species shows definite behavioral modification to different envi
ronments. For example, at the northern extreme of the range, in
southwestern Utah, desert tortoises have a “summer range’’ and a
“winter range.’’ In summer the tortoises wander over the desert
flats, retiring at night to short burrows which they excavate at the
bases of bushes. However, in winter, when temperatures get ex
tremely low, the tortoises repair to the foothills, and during the
nights and cold periods they retire to communal burrows which
may extend as far as 33 feet from mouth to end. These burrows are
enlarged very slightly each year; but to make a burrow a foot
deeper in the hard soil appears to take many years. Since a tor
toise can only escape the extreme winter cold in a burrow at least
fifteen feet long, one immediately wonders how a new hibernation
burrow ever gets started. The answer appears to be that all bur
rows currently in use in this area were already at least fifteen feet
long when the climate changed, perhaps 5,000 years ago, from
moderately warm winters (in which the tortoises were safe even in
shallow burrows) to present very cold winters.
Further to the south, in Arizona, the tortoises also have to
hibernate, but here the winters are sufficiently mild for a tortoise
to be quite safe simply in an enlarged ground squirrel burrow in
the side of the bank of a dried-up stream. Such hibemacula may be
used for several years in a row, but are destroyed during occa
sional wet periods when the streams run again. These hibernation
sites are always on south-facing banks, to take full advantage of all
available solar warmth.
Further south still, in central Sonora, Mexico, it appears that
the tortoises do not burrow at all; the winters are sufficiently mild
to make this habit unnecessary.
The desert tortoise is nomadic in its habits. In Sonora the tor
toises wander great distances along the dry stream beds that dis
sect the area. This tortoise is often active even during the middle
of the day, when temperatures can be extremely high. Mating
takes place in May and is often preceded by protracted combat
between males. Nesting commonly takes place in June, and the
eggs are hard-shelled and either spherical or slightly ovate. Grant
(1936: Zoologica, 21: 225-229) gives the clutch size of five females
as two eggs in one case, five eggs in three cases, and six eggs in
one case. A captive specimen kept by Forrest and Ray Poorman
laid between 6 and 13 eggs a year for six years. A typical egg
measures 1.8” x 1.5”, but there is much variation. Incubation
takes three or four months. The hatchlings, about two inches in
length, are somewhat more elongate than those of many tortoises
and have predominantly very light colored shells with dark scute
419
www.Ebook777.com
-
Leatherback turtle, Dermochelys coriacea, on the nesting beach in French

Guiana.
Head of a juvenile arrau turtle, Podocnemis expansa, a giant side-neck from

the Orinoco and Amazon.
420
421
www.Ebook777.com
borders. In some specimens there are dark blotches on each of the
vertebral and costal scutes. The periphery of the carapace, light
yellow in color, Is markedly scalloped.
The desert tortoise is now protected by law in California,
where it Is illegal to “sell, purchase, needlessly harm, take, or
shoot any projectile at a desert tortoise.” Despite this law, many
Californians keep desert tortoises, and only in relatively few cases
were these tortoises in captivity before the institution of the ban, or
descended from long-term captive specimens, and thus outside the
provisions of the law. In Arizona the species may not be imported
or sold, and each resident is allowed to keep no more than one spe
cimen. Fortunately this species is apparently never used for food.
In southern Texas and northeastern Mexico lives the smallest
living species of North American tortoise, Gopherus berlandieri,
commonly known as Berlandier’s tortoise, or as the Texas tor
toise. This species in Texas occurs as far north as Aransas,
Refugio, Bexar, Medina, and Uvalde Counties, while in Mexico the
range reaches eastern Coahuila, Nuevo Leon, and Tamaulipas at
least as far south as Ciudad Victoria and Soto la Marina, and with
one or two records from near Ciudad Valles and Tampico.
The Texas tortoise is about 6 to 7 inches long when mature; the
record length is 8.5”. It has a short rounded shell with projecting
and usually strongly forked gulars. The interhumeral seam is
longer than the intergular seam. The head is narrow and pointed,
similar to that of G. agassizi. The pygal region is strongly down-
curved, particularly in mature males. The “scent glands” under
the chin are particularly prominent in this species. In color the
carapace is dark brown to black on hatching, with light yellow
scute centers. The pigmented areas become lighter with growth,
young adults having brown shells, still with yellow areolae. In old
specimens the carapace assumes a uniformly light brown or horn
color. The nuchal scute is often lacking in this species, but when
this happens, the first marginal scutes are separated by the front
part of the first central scute. The plastral coloration is variable,
but usually consists of an irregular but more or less symmetrical
arrangement of very dark brown and buff. The head and limbs
tend to be of the same color as the carapace, but there is often a
broad lighter band along the temporal area. The Texas tortoise
carries alternation in width of the lower ends of the pleural bones
to an extreme degree; the lower ends of the fifth pleurals (at least)
are often completely excluded from contact with the peripherals.
Texas tortoises are particularly abundant in the area west of
Brownsville, Texas, where they concentrate in “lomas,” or low
hills, and very rarely visit the low-lying flats between the lomas.
Auffenberg and Weaver found that on certain areas of such lomas,
population densities equivalent to one tortoise per 82 square
422
The Texas tortoise, Gopherus berlandieri, is the smallest North American tor
toise. It is protected in Texas.
meters existed. The front feet of the Texas tortoise are less
adapted for burrowing (i.e. less flattened) than in other members
of the genus, and one indeed finds that winters and nights are suf
ficiently mild throughout most of the range of the species that a
shallow pallet, barely the length of the tortoise’s shell, is sufficient
for protection from the coldest weather. Proper burrows, there
fore, are not made except in a few exposed situations in the
northern part of the range, where the soil is sandy and thus suit
able for burrowing. Even in these areas, the burrows do not extend
for further than three or four feet.
In the wild state, the Texas tortoise feeds mainly upon grass in
the wetter times of the year, and cactus pads, flowers, and fruits
when the grass withers in the dry season. The most commonly
eaten cactus species is Opuntia lindheimeri.
Details of the courtship of Gophertis berlandieri have not been
described. Nesting takes place during the warmer months of the
year (extreme definitely known nesting dates are June 8 and
August 22), with some indication that trial nests may be made as
early as April, and also that some nesting may take place as late
423
www.Ebook777.com
Plastral view of the Amazonian side-neck Podocnemis sextuberculata. The
tubercles after which the species is named are at the outer rear corner of the
pectoral, abdominal, and femoral scutes; they disappear as the turtle grows.
The seldom-seen Madagascan side-neck, Erymnochelys madagascariensis.

Photo by Wade Ferrar.
424
A rarely-encountered zoogeographic enigma, Erymnochelys madagascarien-

sis.
Young Gabon terrapin, Pelusios gabonensis, the most attractively marked of

the African side-necks.
425
www.Ebook777.com
as November. Auffenberg and Weaver (1969: Bull. Fla. State
Mus., vol. 13, 3: 141-203) found that in 73 nests, one egg only was
present in 46 cases, two eggs in 23 cases, and three eggs in one
case. However, it is possible that more eggs, on average, are laid
in some areas. Forrest and Roy Poorman (1971: Int. Turt. Tort.
Soc. J. 1: 14-16) record that a young female berlandieri in cap
tivity in California laid 3-4 eggs at a time. The eggs are elongate
rather than almost spherical as in other members of this genus
and are about two inches long and 1.4 inches in diameter. Under
natural conditions strong preference is shown for certain circum
scribed areas for nesting, these usually being in well-drained
areas, relatively free of surface vegetation, and often in the drip
zones of shrubs. The hatchling is about 1.6” in length.
Male Texas tortoises have the usual concave plastron and a
relatively long hind plastral lobe, with the points of the xiphiplas-
tra sometimes actually touching the edges of the supracaudal
scute. Males are slightly larger, on average, than females and
have slightly narrower shells.
In recent years the Texas tortoise has been exploited in very
large numbers for the pet trade. This trade was regrettable for two
reasons: the tortoises have a low reproductive potential, so even
though locally numerous, they could not tolerate massive annual
losses; also, although the species does a little better in captivity
than the other members of the genus, it is basically not easy to
keep Texas tortoises alive outside of their native habitat, and it is
probable that almost all specimens sold as pets live only a few
months. Legislation has now been passed making it illegal to col
lect, harm, trade in, import, or export these tortoises in the State
of Texas. There is, however, still a considerable pet trade in this
species, the dealers usually asserting that the animals were col
lected in Mexico. Such assertions are of course extremely unlikely,
and even if the animals were collected in Mexico, it is again very
unlikely that they reached their ultimate stateside destinations
without being shipped through Texas.
Another drain on Texas tortoise populations is the accidental
killing of large numbers by automobiles. Fortunately the density
of major highways throughout the range of the species is relatively
low, but even so those populations adjacent to the highways must
undergo heavy attrition.
More distantly related to the desert tortoise is the Florida
gopher tortoise, Gopherus polyphemus. This species is found in
coastal South Carolina, southern Georgia, throughout Florida (ex
cept for the Everglades), extreme southern Alabama and Missis
sippi, and it just enters extreme eastern Louisiana; it is often very
abundant on small islands off the Gulf Coast of Florida. Adult spe
cimens are nine or ten inches long, while the record (for a long-
426
term captive individual) is normally given as 14.2”. However,

Frank Ligas of National Audubon recently obtained a 16”, 29 lb.
male gopher from the Brighton Indian Reservation, Florida. Theo
dore Roosevelt mentions seeing 18” gophers many years ago, but
it is unlikely that more than one in a million would get that big, and
Roosevelt's account is now usually discounted. The largest I have
seen was a 15” specimen from LaBelle, Florida, that lived in cap
tivity for a few months at Godwin’s Gatorland in Kissimmee,
Florida.
The Florida gopher has a very flattened, moderately elongate
carapace, usually of a more or less uniform brown color though
some mottling may be visible, and the areolae are often slightly
darker than the rest of the shell. Hatchlings are very differently
colored, the carapace scutes being of a bright orange-yellow color,
thickly edged with black. Growth annuli are well-developed and
often easy to count, but the shells of old gophers become complete
ly smooth. The nuchal scute is broad and the carapace margin is
slightly scalloped. The sides of the shell are either parallel or
slightly bowed, with their widest point near the middle. The plas
tron is yellowish, wide and flat in shape, with a slight concavity in
males. The gulars are extended somewhat forward and may or
may not be slightly forked. As with all members of the genus, there
is a deep angular notch between the xiphiplastra. The head is
much wider than in agassizi or berlandieri. The scales of the ex
tremities are dark grey-brown, separated by areas of grey in
adults or uniform yellow in most hatchlings. The forelimbs are
markedly flattened and the wrists are almost completely rigid. As
one would expect from such a forelimb structure, the Florida
gopher is a burrower. In fact, deep burrows are dug in all parts of
the range, except for one or two areas on the Gulf Coast of Florida
where the climate is sufficiently warm and moist to obviate the
need for such a construction and the ground unsuitable for digging
one anyway.
Gopher burrows are in fact a very characteristic feature of
sandy areas of the deep south. Typically they are most often seen
in ecotonal areas, and may immediately be recognized by their
semicircular profile. A tortoise starts a burrow soon after it
hatches, or takes over a pre-existing shallow depression in the
ground and deepens and widens it as it grows up; only in excep
tional circumstances is a new burrow started late in life. The bur
row, which may be straight or may curve extensively, reaches a
maximum length of about thirty feet and a maximum depth of
twelve feet. It provides a refuge for the tortoise both at night and
throughout the winter; the end of the burrow not only provides pro
tection from temperature extremes, but also provides the high
level of humidity that this species apparently requires. Conditions
427
www.Ebook777.com
Front view of the African side-neck Pelusios castaneus.
Plastral view of the helmeted turtle, Pelomedusa subrufa, a species

widespread in sub-Saharan Africa.
Two views of a young matamata, Chelus fimbriatus.
429
www.Ebook777.com
outside the burrow are often extremely arid, but gopher tortoises
in captivity that are denied access to burrows usually become
severely dehydrated and eventually die. Very young individuals,
however, may be raised in captivity without providing a burrow.
The gopher tortoise is herbivorous and under natural condi
tions eats large quantities of grass and leaves. Its daily routine
often involves passage along well-defined “grazing trails” that
extend to and from the mouth of the burrow.
The burrow of the gopher tortoise provides a refugium for
many other organisms. Apart from at least 32 species of arthro
pods, rattlesnakes are often found in gopher burrows, as are opos
sums and gopher frogs (Rana capito). Constant maintenance of
the “grazing trails” mentioned above encourages the growth of
certain types of native grass, such as carpet, St. Augustine, and
centipede grass. Since the seeds of these grasses are unaffected by
the digestive process of the tortoise, grazing of these plants by tor
toises is probably of importance in their maintenance and spread.
One often sees ribbons of these grasses winding through seas of tal
ler, coarser grasses or bushes; these appear to be a direct result of
the feeding and defecatory processes of the tortoises.
Gophers appear to be very coy about their copulatory and
nest-building functions, and these processes have found few obser
vers. However, we do know that, when courting, male gophers bob
the head up and down in a fashion very reminiscent of that of a toy
tortoise of the type in which the head and neck are suspended by a
single string. This behavior actually seems to attract females
from some distance. The rate of bobbing increases as the female
gets nearer, and when she comes within reach, the male brings her
to a state of immobility and sexual receptiveness by biting her
forelimbs and the front edge of her shell. It appears likely that the
change in behavior of the male from bobbing to biting is triggered
by the female wiping her chin with her forelimbs; a specialized,
pointed scale on each forelimb apparently removes a small quan
tity of the secretion of the chin glands, and the male may be
responding to the odor of this. The rate of head bobbing as well as
the nature of the secretion appear to differ in the four species of the
genus, and these differences may well have prevented interspeci
fic matings at that time in the Pleistocene when the species were
sympatric.
Patterns of mate-seeking by Gopherus polyphemus have been
studied in recent years by John Douglass, working at the Archbold
Biological Station in Highlands County, Florida. Douglass found
that dominant males maintain their own burrows and visit as many
as four different females at separate burrows during spring, sum
mer, and fall. The male head-bobs at the entrance to the female’s
burrow and awaits her emergence. One female may be visited by
430
three different males during a season, but a male will seek out an
individual female from year to year even if her preferred burrow
changes. Both sexes will fight to defend their burrows, and males
will fight each other over access to females and their burrows. The
largest males are the dominant individuals, but because of their
slowness of movement, their tours of harem inspection take some
time and subordinate males may take advantage of the absence of
the dominant male to mate with preferred females.
The distance a gopher tortoise ventures from its burrow each
day depends upon the density of suitable food plants in the vicinity.
When disturbed within a few yards of the burrow, they make
straight for the entrance with surprising speed and disappear
within. Perhaps correlated with this flight rather than withdrawal
reaction is the very thin shell of the gopher tortoise; in many in
dividuals it is actually fenestrated.
Gophers usually but not always make their nests and lay their
eggs in the mound of sand that they excavate out of their burrow.
Recorded clutch sizes range from four to seven eggs. The eggs are
almost spherical and range in diameter from 1.5” to 1.73”. The
nesting season extends from late April to July.
Gophers are frequently eaten by rural people in the deep south
and appear to be a particular favorite among black people. Human
consumption, frequent road kills, extensive habitat destruction,
and casual taking-home of gophers by people in all walks of life are
causing a steady reduction in total population numbers. At the
present time there are no laws against collecting the species in any
state, although it is now illegal to buy or sell gophers in Florida.
One of the biggest cheloniological surprises of the twentieth
century was the discovery of a very large fourth species of
Gopherus in north-central Mexico. At least two specimens of this
tortoise were collected as long ago as 1918 and deposited in the
United States National Museum, but it was not until 1959 that John
Legler studied all the available specimens and pronounced them
as belonging to a new species, Gopherus flavomarginatus.
Gopherus flavomarginatus is known in English as the giant
Mexican gopher tortoise or as the Bolson tortoise. In Mexico it is
called tortuga grande, or tortuga llanero. It has a presumably
relictual distribution in an internally drained basin in north-cen
tral Mexico, centered around the area where the States of Chihua
hua, Coahuila, and Durango come together. Definite locality
records exist for the vicinity of Ceballos, Durango; Americanos,
Coahuila; and Carrillo, Chihuahua.
The Bolson tortoise appears to be more closely related to
Gopherus polyphemus than to G. berlandieri, even though the
closest part of the range of berlandieri is only 100 miles away,
431
www.Ebook777.com
An adult matamata turtle, Chelus fimbriatus, in a zoo pool. Photo by Dr.
Herbert R. Axelrod.
Plastral view of a juvenile South American side-neck, Phrynops gibba, from

Surinam.
432
www.Ebook777.com
Detail of head of a young Phrynops gibba from the Upper Commewijne River, Surinam.
while polyphemus is found nowhere nearer than extreme eastern
Louisiana. In fact, as long ago as 1888, Duges (La Naturaleza, 1
(ser. 2): 146-147) collected Gopherus in Chihuahua, which he con
sidered to be referable to polyphemus. G. flavomarginatus differs
from polyphemus in a number of characters: the carapace of
adults is flared posteriorly, reaching its widest point opposite the
anterior parts of the fourth costal scutes; the carapace is lighter in
color, especially toward the middle of the marginal series; the
extremities are also lighter; the intergular seam is always longer
than the interhumeral seam; and the adult size is considerably
larger.
Gopherus flavomarginatus has a pale yellow or straw-colored
carapace when young, becoming pale brown in adults. The areolae
of the carapace scutes are darkened, and the middle marginal
scutes have particularly dark areolae on a light yellow, strongly
contrasting ground color. The soft parts range in color from dirty
cream to yellow; the largest scales on the limbs and head have
dark centers. The carapace is very similar in shape to that of poly
phemus, except for the slight flaring above the hind feet. The head
is relatively larger than in any other member of the genus. The
The Bolson tortoise, Gopherus flavomarginatus, of north-central Mexico is

probably a relict of a wider Pleistocene distribution.
head scales are usually irregular, but ordinarily there are two pre-
frontals, one frontal, and two parietals. Both sexes have paired
chin glands and also have a pair of yellowish, elongate, black-
tipped spurs on each thigh.
The courtship of flavomarginatus is known to involve periods
of head bobbing, but no other details are available. Legler was in
formed by local people in Durango that the young are commonest
in late summer and early spring, but it is doubtful that this obser
vation has any bearing on the hatching season. From measure
ments of the plastral areolae of adult specimens, it seems that
hatchlings have a plastral length of about two inches. There is a
high degree of scute abnormality in this species, many specimens
having extra central and costal scutes.
Just how much larger flavomarginatus is than polyphemus is
open to question. The largest individual available to Legler had a
carapace length of 14.6 inches, only just larger than what was until
recently the largest polyphemus (14.1 inches). However, the flavo
marginatus record was based on a series of only fifteen individuals
while the 14.1” gopher was the largest of thousands. An advertise
ment in the May-June 1967 issue of the International Turtle and
Tortoise Society Journal offered for sale flavomarginatus measur
ing up to 17” in length, and stated that the species is “recognized
as reaching a 26” size.” My friend Bruce Deuley saw the shell of a
23” specimen in the town of Ceballos, Durango, and Auffenberg
informed me that, judging by the size of certain burrows, a cara
pace length of about 27” is reached. Local people stress that the
largest specimens measure nearly a meter in length. This appears
to be a species that, like Geochelone denticulata, matures at a rel
atively modest size (about one foot), but which may on occasion
become very much bigger.
It is probable that at the end of the Pleistocene the Bolson tor
toise reached all the way from southern New Mexico to southern
Zacatecas; but its steadily shrinking range is now only about 75
miles long. These tortoises live in burrows which may be eight feet
or more in length, while Ray Pawley found a burrow which
branched every four feet or so, with a total of 22 feet of tunnel work
which reached a maximum vertical depth of over six feet. The bur
rows appear to occur in definite clusters rather than at random
and are usually found at the edges of large flat areas, almost bare
of vegetation, which almost certainly represent the floors of dried-
up lakes.
The habits of this species are poorly known, but it appears that
the Bolson tortoise, like polyphemus, grazes along feeding paths
cut through the grasses surrounding its burrow. It ventures out in
the early morning and late afternoon hours, but avoids the
extreme cold of night and heat of day by staying deep underground
at these times.
435
www.Ebook777.com
Two views of Phrynops rufipes, an extremely rare side-neck from the Upper
Amazon.
436
Plastral view of Phrynops hilarii, a large side-necked turtle from southeastern

South America.
Phrynops geoffroanus geoffroanus photographed at the Frankfurt Zoo.
www.Ebook777.com
The long-drawn-out natural extinction of Gopherus flavomar
ginatus, already at a very advanced stage, has in recent years
accelerated greatly as man has invaded the area and killed the
tortoises for food, established irrigation projects, and planted cot
ton fields. Bruce Deuley even reports that the tortoises are killed
wantonly simply because they compete with cattle for the sparse
grass. On the other hand, Ray Pawley found that the local people
actually put out food for the tortoises in some areas and refrain
from taking more tortoises than the populations are thought able
to stand.
GENUS PSAMMOBATES
A pattern of radial streaks on each of the carapace scutes
turns up repeatedly in the Testudinidae; the so-called “starred”
tortoises of Asia and Madagascar have been described earlier in
this chapter. The present genus, Psammobates, includes three
species of South African starred tortoises. This genus is set apart
from Geochelone by the retention of the primitive hexagonal shape
of the neural bones and by the lack of the advanced pygal pattern
shown by Geochelone (in Psammobates the pygal bones number
one or two; if two they are separated by a straight transverse
suture). However, the skull may be considered advanced in that it
has lost the maxillary ridging; the longer-than-broad shape of the
gular scutes is also an advanced feature. The three species of
Psammobates are all very small tortoises (less than six inches
long); they never develop hinged shells. Two of the three species
(geometricus and oculifer) are of relatively constant appearance
and are not separated into subspecies. The third species, tentorius,
is so variable that literally dozens of names have been given to dif
ferent color phases or local variants. Only three of these are now
considered valid, but the species nonetheless shows a bewildering
range of variation, and extreme specimens of, say, Psammobates
tentorius verroxii, are fully as different as extreme variants of
that epitome of variability, the eastern box turtle (Terrapene Caro-
lina Carolina). It is only because the variants show poor or no geo-
graphic correlation that they are not considered valid subspecies.
The geometric tortoise, Psammobates geometricus, is the lar-
gest of the genus, the present-day maximum length appearing to
be in the vicinity of six inches, with old and unconfirmed records of
specimens up to ten inches long. This is a handsomely marked spe-
cies with a highly convex carapace, often with distinctly swollen
vertebral scutes. Each of the carapace scutes has a light yellow
areola with radiating yellow streaks. These number 8 to 15 on the
vertebrals, 9 to 12 on the costals, and 2 to 4 on the marginals. The
hind margin of the carapace is slightly reverted and the edge is
somewhat scalloped—feebly serrated in some individuals. The
438
plastron is anteriorly truncate, with a shallow nick between the

gular scutes. A single axillary scute is present on each side; the
inguinal is moderately large and makes contact with the femoral
scute. The plastral pattern involves a pattern of light areolar areas
with dark scute borders; the dark borders are usually penetrated
by light radiating streaks, but nevertheless the dark scute borders
are a useful identification feature. The prefrontal and frontal
scales on the head are fragmented. The beak is at least moderately
hooked. The forelimb is anteriorly covered with a few large, un
equal, scattered tubercular scales. There are no tubercles or spurs
on the thighs or tail.
As is usual among starred tortoises, the hatchling presents a
very different appearance from the adult. Many hatchlings of geo
metricus have a characteristic pattern technically known as the
“luteola” phase, in which each of the vertebrals and costals is
marked with a single broad yellow “X”. The plastron at this stage
is mainly black, but with a yellow margin and a yellow line along
the midline which sends out lateral expansions on the humeral and
abdominal scutes.
The geometric tortoise has the unfortunate habit of living in a
type of habitat known as “suurveld,” a mixture of evergreen
shrubs and grass growing on a very acid, sandy substrate. Such
habitat is fast disappearing in South Africa, where it is being
ploughed up for crop cultivation, and with it, the geometric tor
toise has receded almost to the point of extinction. The species only
survives in a few tiny areas of more or less virgin habitat in Cape
Province, in the vicinity of the towns of Paarl, Malmesbury, Dar
ling, and Ceres. The species has been recorded as far north as
Endekuil, but it is not known if it still occurs there. A hazard to the
future of geometricus in the Ceres Valley is an irrigation canal
which bisects the valley; Mr. Reinhold Rau of the South African
Museum has found several drowned geometricus in this canal.
However, it has been established that this species can maintain a
population even in a few acres of virgin habitat entirely sur
rounded by farms, and the IUCN (International Union for the Con
servation of Nature) has now purchased five acres of such an area
near Paarl, which will be fenced and maintained in perpetuity.
James Juvik made the interesting observation that female
geometricus appear to be much commoner than males. This obser
vation is confirmed by the studies of Mr. Rau, and it appears to be
a real phenomenon rather than just sampling error or an indica
tion of an accidental collecting bias.
The African serrated tortoise, Psammobates oculifer, is found
in inland grassland savannas of extreme western Transvaal and
western Orange Free State northwest through Botswana and
South-West Africa almost to the frontier of Angola. This species is
www.Ebook777.com
Toad-headed turtle, Phrynops nasuta nasuta, from Surinam.
Plastral view of young toad-headed turtles, Phrynops nasuta nasuta, from

near Berlein, Surinam.
440
441
www.Ebook777.com
recognized by its very serrate carapace margin; even in adults,
both anterior and posterior carapace margins are usually strongly
serrate. The carapace is not as elevated as that of geometricus. A
posteriorly-expanded nuchal scute is present. The vertebral and
costal scutes are slightly swollen but are not conical. The ground
color of the carapace is brownish-yellow with a very variable
number of dark brown or black radiating streaks; typically these
number from six to ten on each of the vertebrals and costals. In
many individuals the black streaks are distally expanded to the
extent that the scute would better be described as black with a
light areola and light radiations. The pattern is continued on the
plastron, though the dark streaks are not as dark as on the cara
pace. The plastron is openly notched both anteriorly and posterior
ly. The axillaries are not divided, and each inguinal is in contact
with the femoral. In some individuals the pectorals fail to meet on
the midline.
The beak of oculifer is hooked and tricuspid, and the jaw sur
faces are more or less serrate. The frontal and prefrontal scales
are fragmented. The forelimb is faced with a few unequal tuber
cular scales, one of which is conspicuously enlarged. There is at
least one enlarged subconical tubercle on each thigh, often sur
rounded by smaller subsidiary tubercles. There may or may not be
tubercles on the heel.
442
The habits of the serrated tortoise have not been studied in any
detail. Fitzsimons, quoted by Loveridge and Williams (1957),
watched the courtship of a pair of oculifer on a road near Kimberly
on November 29; the male reportedly scuttled around the female,
butted against her shell, and periodically emitted short, low,
grunting coughs. A typical egg measured 1.55” x 1.22”.
The maximum carapace length of the serrated tortoise
appears to be a little over 5 inches. In captivity, the species is
reported to be very delicate, requiring very high temperatures and
low humidity.
The African tent tortoise, Psammobates tentorius, is widely
distributed in South and South-West Africa, and shows such a
bewildering degree of variation (sometimes geographically corre
lated and sometimes not) that a completely satisfactory allocation
of subspecific names to the various distinct populations has not yet
been made. Loveridge and Williams reduce the recognizable sub
species to three (tentorius, trimeni, and verroxii), each of these
showing a wide range of variation, but nevertheless showing cer
tain distinctive characters.
Psammobates tentorius tentorius is a small starred tortoise
from southeast Cape Province, the distribution being said to be
mostly south of the 32nd parallel and to be concentrated in the
region from Grahamstown to Uniondale. The carapace is convex,
with rounded sides, and in most individuals the dorsal scutes,
especially the vertebrals, are markedly conical. The nuchal is
usually small and may be completely absent. The carapace is yel
low or orange in color, with 8 to 12 black radiations on each verte
bral, 12 to 14 on each costal, and 3 to 4 on each marginal. The
plastron is anteriorly truncate, with two or more axillary scutes on
each side and one or two supplementary inguinal scutes. The mar
gins of the plastron are orange or yellow, with a large central dark
figure covering most of the underside of the animal. The plastral
figure may be solid black or it may show a tendency to be more in
tense along the seams.
The beak of tentorius is hooked and the jaw surfaces are ser
rated. The dorsal head scales are extensively fragmented. There
may or may not be a large, subconical tubercle on each thigh;
there are no tubercles on the heel. The southern tent tortoise, as
this race is called, reaches a carapace length of about 5.5 inches.
Reproduction is said to take place in September, 2 to 3 eggs being
laid, which measure, on average, about 1.2” x 0.9”.
Psammobates tentorius trimeni, the western tent tortoise, is
found in extreme western Cape Province, from Lambert’s Bay
north to the Orange River, and across the river into Great Nama-
qualand. This race is distinguished from tentorius by the smaller
number of dark rays on the carapace scutes (only 4 to 8 on each
costal and vertebral, the dark rays being distinctly broader than
443
www.Ebook777.com
Broad-shelled river turtle, Chelodina expansa, from southeastern Australia.
A juvenile saw-shelled turtle, Elseya latisternum, from northeastern

Australia.
444
www.Ebook777.com
Common long-necked turtle, Chelodina longicollis, from Australia.
The fewer and broader dark rays on the scutes identify Psammobates ten-
torius trimeni. Photo by Dr. Robert Mertens.
the intervening yellow or orange bars) and by the invasion of the

central dark figure on the plastron by radiations or incursions of
yellow. The largest specimen on record (a female) had a carapace
length of 4.8”. An egg measured 1.36” x 0.95”. Males have longer
tails than females but lack the patch of enlarged scales on each
side of the tail. The males also have more protuberant foreheads
than females, and the sides of the anterior and posterior plastral
lobes are less convergent.
In the arid bush country inhabited by trimeni, standing water
is a rarity, and the tortoise has evolved an extraordinary method
of taking advantage of light rainfall to satisfy its water require
ments. When rain falls upon a thirsty tent tortoise, it adopts a
peculiar stance with the posterior part of the shell raised on the
extended hind limbs, while the head is extended between the out
wardly turned, fully extended forelimbs. Rain falling on the cara
pace is directed forward and downward but is prevented from fal
ling off the sides of the shell by the reverted marginal scutes. In
stead the water flows off the shell on each side of the nuchal and is
directed toward the mouth by the outstretched forelimbs. This
446
method of obtaining drinking water is also shown by some other

tortoises with recurved marginal scutes, including Kimxys
homeana and Homopus areolatus.
Psammobates tentorius verroxii, the northern tent tortoise, is
found in northern Cape Province from Burghersdorp east to Willis-
ton and northwest to O’okiep, and in South-West Africa from Great
Plastral view of
Psammobates ten
torius tentorius
showing the dark
area without yel
low markings.
Photo by
John Visser.
Namaqualand to Rehoboth. This race lacks the conical scutes usu

ally present in tentorius and trimeni, and the plastral pattern, if
present at all, is not sharply defined. Many specimens show a
great reduction in the dorsal radiations, either in intensity of con
trast or in number, or both, and in some specimens (such as in the
color form “bergeri,” named after an invalid subspecies of that
name), there are no radiating streaks at all.
It appears probable that verroxii only lays a single egg each
year, in or around the month of December. The diet is known to
include the leaves of the Mesembryanthemum bushes under which
the tortoises are wont to hide.
447
www.Ebook777.com
A New Guinea side-neck turtle, Elseya novaeguineae.
A very rare and localized side-neck, Pseudemydura umbrina, from Western

Australia. Photo by Dr. Robert Mertens.
448
GENUS HOMOPUS
The genus Homopus includes four small tortoises restricted to
South Africa. The genus is defined on the basis of the combination
of a relatively primitive pattern of neural bones (i.e. including
hexagonal and quadrilateral elements, but none octagonal) with
an advanced pygal pattern (i.e. an enlarged suprapygal enclosing
a smaller one between its ventral rami, as in Geochelone and
Gopherus). The triturating surfaces of the maxilla are unridged,
and the gular region is somewhat thickened and produced. Both of
these are advanced conditions, but the short, broad gular scutes
are a primitive feature.
The four species of Homopus can be aligned into two species
groups: signatus and boulengeri, characterized by fusion of the
distal carpal bones, the presence of five claws on the front feet,
and the presence of twelve marginal scutes on each side; and areo-
latus and femoralis, in which the distal carpals remain unfused,
and which have only four claws on each front foot, and only eleven
marginals on each side, as in most tortoises. Hewitt in 1931 pro
posed a new genus (Pseudomopus) for the first two species, but
acceptance of this would obscure the demonstrable naturalness of
the genus Homopus. In any cause, a much older name (Chersobius
Fitzinger 1835) is available if separation of signatus and boulen
geri is ever considered justified.
The speckled tortoise, Homopus signatus, is a very attractive
little tortoise from western Cape Province (mainly Little Nama-
qualand) northward into southern South-West Africa. It is one of
the smallest living species, not being known to exceed a carapace
length of four inches. The shell is rather flattened and is sharply
declivous posteriorly. The carapace margin is serrated and some
what expanded and reverted. The scutes of the carapace are con
centrically striated and have slightly recessed areolae and seams.
The ground color of the carapace is white, yellow, orange-yellow,
or yellow-green, but is always heavily overlaid with black speck
ling which may form interrupted radiating streaks. The plastron is
anteriorly truncated, with the comers of the gulars slightly pro
jecting. The carapace pattern is continued on the plastron, though
the black speckling is replaced by brown. The head is basically
yellowish on the top and sides, darker around the jaws, and with
black spots on the light areas. The beak is at best weakly hooked.
The forelimbs are covered with very large, overlapping scales.
The habits of the speckled tortoise have not been studied. It is
said to be fairly numerous on granite kopjes 16 miles from Bitter-
fontein and to reach south as far as Clanwilliam Mountain.
Homopus boulengeri, though closely related to H. signatus,
completely lacks the attractive markings of the latter species and
has a uniformly brown, or occasionally olive-green, carapace.
449
www.Ebook777.com
450
A speckled tortoise, Homopus signatus, from Little Namaqualand, South

Africa. Photo by J. Visser.
This is a slightly larger species, reaching a carapace length of

nearly 4.5 inches; the shell is depressed, with expanded but not
reverted margins, and with at most feeble marginal serrations.
The plastron is similar in shape to that of signatus, anteriorly
truncated and projecting slightly at the corners of the gular scutes.
Single axillary and inguinal scutes are present on each side. The
naked parts of the skin in living specimens are said to be bright
yellow with minute orange scales. In preserved material the jaws
appear brown and the head and limbs light olive-green.
The range of the donner-weer tortoise, as this species is locally
called, is restricted to the Karroo Plain area of south central Cape
Province, with the exception of an apparently reliable record of a
pair having been collected from Plateau Farm, near Aus, South-
West Africa, hundreds of miles away from the Karroo Plain.
The apparently single egg of Homopus boulengeri is relatively
enormous; one measured by Siebenrock, taken from the body of a
female tortoise, measured 1.53” x 0.87”, its length about 40% of the
carapace length of the average female. Passage of such an egg
through the rather narrow posterior shell opening necessarily in
volves some kind of spontaneous mobility of the hind plastral lobe.
It is possible that the calcium for the eggshell is partially derived
451
www.Ebook777.com
from a de-ossification of the anterior part of the xiphiplastra, so
that by the time the egg is hard-shelled, the hind plastral lobe is
somewhat mobile.
Perhaps the best-known species of Homopus is the parrot-
beaked tortoise, Homopus areolatus, a lowland form found along
the coastal regions of Cape Province south of latitude 32°S. This is
another small species, not exceeding a carapace length of 4.5
inches. The carapace is moderately domed but somewhat flat
tened dorsally. The carapace margin is virtually unserrated, is not
expanded, and may be slightly reverted posteriorly. The areolae
are slightly inset, the scutes themselves being slightly swollen and
with well-marked growth annuli. The nuchal is variable in shape,
being elongate, subtriangular, or quadrate. In color the carapace
is yellowish-olive to greenish, each scute with a red-brown center
and with dark margins. The plastron is basically yellowish anteri
orly with a brownish area along the midline in most specimens and
with axillary and inguinal scutes divided so that they number up to
four on each side. The head and limbs are of a light buff color. The
beak is strongly hooked and tricuspid; the jaw edges are finely
serrate. The forelimbs are covered with strikingly large overlap
ping pointed scales. The claws are long and sharp. Some speci
mens have an inconspicuous tubercle on the hind side of each
thigh. There are no spur-like tubercles on the heels. The plastron is
virtually flat in both sexes; by contrast, males of boulengeri have
marked plastral concavities.
According to Hewitt, the parrot-beaked tortoise is a shade-
loving form which inhabits sour-grass areas from sea level to the
summit of the Zuurberg. When picked up, these tortoises struggle
violently rather than retire into their shells; this may be a more
effective response when the animal is grabbed by a large preda
tory bird—secretary birds are known to eat tortoises of this spe
cies. Again according to Hewitt, the nest holes are about three
inches deep and the eggs, 2 to 5 in number but usually three, ave
rage about 0.87” x 1.18” in size. Rose found that incubation takes 7
to 8 months.
The Karroo tortoise, Homopus femoralis, inhabits grassveld
up to 5,500 feet in the eastern Karroo in Cape Province and also
western Orange Free State. It is the largest member of the genus,
measuring up to 6.2” in carapace length. This species is closely
related to areolatus, but the carapace scutes are not swollen and
are more uniform in color, being olive or brown, sometimes with a
dull red or orange tinge; in the young each scute is narrowly edged
with black. The plastron is greenish or yellowish, each scute with a
blackish-brown marking anteriorly in the young. The scales of the
extremities are yellow or pale brownish, with a salmon or orange
tinge to the naked areas. The axillary scute of the bridge is single,
452
Anterior views of Homopus species.
Homopus boulengeri,
female.
Homopus areolatus.
453
www.Ebook777.com
but the inguinals number two or three. The beak is only slightly
hooked. In contrast to areolatus, there is a very large conical
tubercle on the rear of each thigh and there are also large, conical,
spur-like tubercles on each heel. According to Hewitt, the eggs are
quite soft when deposited and measure about 1” x 1.2”.
GENUS KINIXYS
The African genus Kinixys, by virtue of its unique dorsal
hinge, is one of the most distinctive genera of the Testudinidae.
The hinge is situated between the fourth and fifth pleural bones
and between the seventh and eighth peripheral bones. This posi
tion corresponds to the junction of the second and third costal
scutes and the seventh and eighth marginals. The hinge allows the
posterior part of the carapace to be lowered when the tortoise is
molested, presumably offering some degree of protection to the
hind feet and tail region. The hinging may also assist to some
degree with respiration and oviposition.
Other generic characters of Kinixys include the hexagonal
neural bones and the anterior extension of the front peripheral
bones and marginal scutes. The hinge is not present in young spe
cimens, and these can easily lead to confusion. Indeed, the type
specimen of Kinixys belliana nogueyi, a juvenile without any sign
of a hinge, was originally placed in the genus Homopus. The hinge
gradually develops, starting just behind the bridge with a break
down in the sutural connection between the seventh and eighth
marginals and gradually migrating dorsally. In fully mature Kin
ixys there is usually a broad band of flexible connective tissue
extending to the upper part of the fourth and fifth pleural bones.
The hinging action involves some flexure of the third central
scute; the fourth neural bone, which lies under this scute, remains
in loose sutural connection with its neighbors. In some few adult
specimens the hinge completely fails to develop.
Juvenile Kinixys lacking the dorsal hinge may be identified by
the relative proportions of the costal scutes; the lower margin of
each third costal is foreshortened so that it is shorter than the
lower margin of the fourth costal.
Other unusual features of this genus include the virtual non
development of the usual alternation in width of the lower ends of
the pleural bones and in the non-coincidence of the costomarginal
seam and the pleuroperipheral suture.
Two of the species of Kinixys, K. erosa and K. homeana, are
quite similar in many respects and occupy much of the same range
in West and Central Africa. This apparent defiance of the normal
ecological rules, paralleled by certain other reptiles (such as the
two species of dwarf caiman in South America) is still without
454
explanation and will remain so until the natural history of these

poorly-known species has been properly studied.
Schweigger’s hinge-back tortoise, Kinixys erosa, is found from
Uganda and the Congo northwest to the Gambia, there being
records from Cabinda, Gabon, Rio Muni, Cameroon, Ghana, Ivory
Coast, Liberia, and Sierra Leone. This is the largest species of the
genus, reaching a maximum carapace length of about thirteen
inches. The carapace, either uniform brown in color or with black
areas surrounding the areolae, is somewhat flattened and elon
gate, with a strongly scalloped margin and strikingly flared
anterior marginals. The plastron is typically yellow, with very
large brown or black areas on each of the plastral scutes (except,
in most cases, for the gulars). The plastron is thickened and pro
duced anteriorly, with projecting and slightly forked gulars. Three
or four axillary scutes are present on each side, and a single in
guinal, which makes contact with the femoral. As in all members
of this genus, the hind lobe of the plastron is short and rounded and
only shows a very shallow, broad, median notch. The marginal
scutes usually number eleven on each side, and the nuchal scute is
almost always absent.
The head of erosa is yellow, marked with brown. The beak is
hooked and shows a single cusp. The jaw edges are not dentate,
and the scales on top of the head are extensively fragmented. The
front faces of the forelimbs are adorned with several greatly
enlarged pointed scales. The tail of the male is longer than that of
the female and is equipped with a terminal homy spur. Males also
have slightly concave plastra.
The habitat of Schweigger’s hinged tortoise is said to embrace
moist rain forest, marshes, and the’shady banks of rivers. It tends
to remairr concealed for much of the time beneath dead or living
vegetation, digging itself into vegetable debris using the forked
gular projection. There is some evidence that the tortoise some
times voluntarily takes to water and even that they swim and dive
for food. Captive specimens are omnivorous in diet.
Copulation has been observed by K. Blackwell in captivity in
Nigeria. Blackwell’s account of a mating attempt on April 9 is as
follows:
“The male was observed to approach the female from
the side and try to mount her. After several unsuccessful at
tempts he succeeded in climbing upon her from the tail end.
The female at this time was lying with her plastron parallel
to the ground and her tail extended beyond the edge of the
shell, also parallel to the ground.
“The male raised himself with his forelegs, pushing on
the female’s carapace so that his shell was at an angle of
almost 90° to the ground. His tail being angled to his body
455
www.Ebook777.com
and placed in a forward position between his fully extended
hind legs, thus lying under and slightly to the left of the fe
male’s tail. At this stage the two cloacas touched and united.
The male then lowered his forelegs so that he again rested
on the female’s shell. The female extended her hind legs so
that her body was angled at about 15° to the ground. The
male then extended his neck fully in a downward curve
towards the female’s neck with his mouth wide open. The
female responded by extending her neck upwards and back
wards towards the male’s open mouth. It seemed that at this
stage the male would grasp the female’s neck in his mouth;
this did not occur, however.
“At the moment when, presumably, orgasm occurred
the male emitted a series of hissing squeaks. Soon after this
the female began to walk in a circle, the male still united,
being dragged along until he finally slid off her back. She
then retreated into a comer and withdrew into her shell. ”
Four eggs laid by a captive tortoise had a slightly flattened
ovoid shape, measuring 1.22” x 1.42” x 1.57”. The young differ
from the adults not only in lacking the hinge, but also in having a
strongly spinose carapace margin, each of the marginal scutes
forming at least one spine, and the rearmost marginals and the
supracaudal having three or four spines on each. A Nigerian
hatchling measured by Blackwell was 40 mm long and 40 mm
wide.
Home’s hinged tortoise, Kinixys homeana, is known from the
Congo, Rio Muni, Cameroon, Nigeria, Ghana, Ivory Coast, and
Liberia. In many features this species is similar to K. erosa, but
there are several constant differences. The most obvious peculi
arity of homeana is the peculiar shape of the rear part of the cara
pace ; the shell comes to a strong point at the front of the fifth ver
tebral scute, the region behind this being completely vertical or
even overhanging. In the other species of Kinixys the hind part of
the shell may slope down sharply, but never from the anterior part
of the last vertebral. Home’s hinged tortoise also has a nuchal
scute (except in occasional variant specimens), and there are
more enlarged scales on the forelimb than in erosa (3 to 5 from the
elbow to outer claw in erosa, 5 to 8 in homeana). K. homeana. also
differs from erosa in the longer and thinner postorbital bar and is
said to be a smaller species, the greatest carapace length men
tioned in the literature being only 8.3”. I foolishly failed to
measure some large specimens of homeana that I once photo
graphed at the San Diego Zoo, but I believe they were well over
eight inches long.
Eggs of homeana measured by A. Villiers were 1.81” long,
1.34” to 1.38” wide, and 1.26” to 1.30” thick. Auffenberg has recent-
456
457
www.Ebook777.com
ly described the peculiar drinking stance of this tortoise, which is
paralleled by certain other African tortoises of the genus Psam-
mobates. When rain is falling and the animal is thirsty, it fully ex
tends its rather long hind limbs so that the hind part of the shell is
well elevated. The forelimbs are bent sharply at the elbows, so that
the distal sections are more or less vertical. Water falling down the
shell is conducted along the “gutters” formed by the reverted shell
margins, while the arrangement of the forearm, wrist, and foot,
assures that the water that spills off the front of the carapace is
conducted to the mouth.
Apart from such isolated observations, the habits of homeana
are not well known. One would expect that interesting courtship
rituals exist to prevent interspecific matings between homeana
and erosa, and it is possible that the strikingly different shape of
the hind part of the carapace of the two species is involved in spe
cies recognition or even in prevention of cross-mating; but such
remarks are purely speculative at present.
All other African hinge-back tortoises are currently referred
to the species Kinixys belliana, a very widespread form found in
rather arid habitats from Senegal to Natal. Only two subspecies
are recognized by most authorities (the four-clawed nogueyi and
the five-clawed belliana), but this is an extremely variable spe
cies, and most of the numerous species or subspecies that have
been described, while appearing very distinctive, must be con
sidered simply as phases rather than as subspecies since they
show little or no geographic correlation.
Bell’s hinged tortoise is known from the Sudan, Eritrea, Ethio
pia, Somalia, Uganda, Kenya, Tanzania, Malawi, Zambia, Rhode
sia, Angola, Botswana, Zaire, Cameroon, and the Provinces of
Transvaal and Natal in South Africa. The race has also been recor
ded from several localities in Madagascar (Amboasary, Nosy-be,
Ambanja). A specimen from an unspecified locality in the
Malagasy Republic has recently been declared the holotype of
Madakinixys domerguei Vuillemin 1972. In fact, apart from a pos
sible slight abnormality of the cervico-dorsal vertebral articula
tion, the specimen is a perfectly typical Kinixys belliana. The sub
species nogueyi is known from Senegal, Cameroon, Nigeria,
Dahomey, Togo, Ghana, Sierra Leone, Guinea, Portuguese
Guinea, the Gambia, and the Central African Republic. This is the
smallest species of the genus, reaching an adult length of not much
more than about eight inches.
K. belliana is distinct from the other two members of the genus
in numerous ways; the carapace pattern, while exceedingly vari
able, is usually radiate, while the carapace margin is neither
spinose nor serrate, being either continuously curved or very
slightly wavy. The rear end of the carapace slopes down steeply
458
from the middle of the fifth vertebral (not from the anterior part of
this scute); there are 5 to 9 scutes on the forelimb from the elbow
to the outer claw; the postorbital bar in the skull is either very thin
and loose or is entirely absent; and the typical habitat is savanna
rather than the virgin forests frequented by erosa and homeana.
The carapace of belliana varies tremendously in height but
usually has a rather smooth lateral profile, lacking the angularity
of erosa and homeana. The front lobe of the plastron is very thick
and is truncated, projecting little if at all beyond the front margin
of the carapace. Two to four axillary scutes are present on each
side, and a single large inguinal. The plastron is posteriorly short
ened and rounded and is only very shallowly notched. The shell
coloration of belliana is described by Loveridge and Williams
(1957) as follows:
‘‘Carapace straw-colored, yellowish-brown, olivaceous

brown or reddish brown; in young uniform, or the areolae
deep brown surrounded by a zone of yellow which may ex
tend to the margins of each shield or be interrupted by fine
black radiations from the dark areolae or be entirely re
placed by black edging. The juvenile coloring may persist in
adults, though usually with the modification that the black
Variation in Kinixys belliana belliana. Photo by G. Hughes.
459
www.Ebook777.com
Kinixys belliana nogueyi is found in central and western Africa.
edging of the shields is irregularly or symmetrically broken

up by yellow pigment, or only the black areolae of the juve
nile pattern persists, in which event it is frequently inter
rupted by a median, longitudinal, yellow streak. In a few in
dividuals, chiefly males, the coloring may be uniform. Plas
tron horn color, uniform or smudged with black, the latter
sometimes in the form of radiations from around the areo
lae.”
Bell’s hinged tortoise appears to become active with the onset

of rains and to remain quiescent or even buried during dry periods.
It has been observed feeding on fruit and fungi and also on giant
land snails (Achatina). It is not known if oviposition is seasonal,
but this may well take place during the wet season. Clutches are
small, probably only numbering between one and three eggs
(three were found in each of two females from Mafia Island by Sie-
benrock). The eggs are elongate and bear some relation to the size
of the female tortoise; a 168 mm tortoise contained eggs 41 x 33
mm, while those of a 192 mm tortoise were 45 x 36 mm.
When picked up, these tortoises show as much variation in
temperament as do common box turtles; some will clamp down
the rear part of the carapace with a hiss and retract all extremities
460
tightly, while others will struggle, snap, and expel noxious fluid
from the cloaca.
Even though only two subspecies of K. belliana are considered
valid by Loveridge and Williams, numerous varieties have been
named and are considered by some even to rank as full species. K.
belliana mertensi was described by Laurent in 1956 from material
from northeastern Zaire, and the range was later extended into
Uganda. However, the race was based on such feeble average dif
ferences in certain plastral proportions that it is unlikely that the
population merits formal trinomial designation. W.H. Archer, who
has observed tortoises in South Africa for over forty years, con
siders there to be no fewer than six species of Kinixys in South
Africa alone (K. darlingi from Mashonaland, Bulawayo, Water-
poort, Duiwelskloof, and Tzaneen; K. australis from the Water-
poort, Mannamead, and Korrenkoppjes areas; K. lobatsiana from
the Onderstepoort/Pretoria District; K. natalensis from Mtuba-
tuba, the Umfolozi Game Reserve, and near the mouth of the St.
Lucia; and K. zombensis from Komatipoort). The various “spe
cies” are reportedly distinguished on the basis of the coloration
and relative height of the carapace. None of these is currently con
sidered valid by most authors, but it is possible that further inves
tigations will confirm the validity of at least some of them, though
only as subspecies of K. belliana.
Still other described “species” that are ignored by most
authors are: K. shoensis from Ethiopia; K. spekii from Tanzania;
K. jordani from Zambia; and K. youngi from Malawi. It seems
likely that those specimens with deeper shell are from open
savanna, while those with depressed shells live in rocky terrain,
where they may seek shelter under boulders.
GENUS CHERSINE
Somewhat related to Kinixys is the bowsprit tortoise, Chersine
angulata. This species is abundant in the more coastal parts of
Cape Province, South Africa, and is common even on Table Moun
tain just beside Cape Town. The species extends north of the
Orange River into Great and Little Namaqualand in South-West
Africa. This species is adult at six or seven inches but has been
known to reach 10.4”; males get bigger than females. This species
resembles Kinixys in having the anterior marginals elongate and
in having the gular region thickened and projecting; on the other
hand, there is no hinge in the carapace, and the neural bones show
the usual tortoise pattern of alternating quadrilaterals and octa
gons.
The carapace of Chersine is distinctively marked with black
triangular markings on the marginal scutes and with a thick solid
black border to each of the central and costal scutes. The ground
461
www.Ebook777.com
color is straw, sometimes with a slight orange tinge. An occasional
ray of light coloration may penetrate one of the black scute bor
ders. The nuchal scute is long and narrow, and there is a slight in
dentation in the carapace margin above the head. The anterior
carapace opening is very small and presumably offers good pro
tection against predators. The plastron is mainly remarkable for
the single gular scute which, particularly in males, covers a mas
sive and greatly projecting epiplastral battering ram. Much of the
area of the plastron from the humerals to the anal scutes is cover
ed with a large symmetrical black figure with somewhat angular
margins. Often this is an unbroken sooty black, but in some speci
mens the intensity of pigmentation is less and the plastral figure
may be interrupted. Each of the under-marginals posterior to the
inguinal has a black marking. The rest of the plastron and the
undersides of the marginal scutes may be either yellow or pinkish-
red ; there is some evidence that there is a geographical correla
tion between the two types of plastral coloration. The head is
small and has extensive black markings, but the chin area may be
suffused with pink. The limbs are black on their dorsal surfaces,
straw-colored below. In captivity, male bowsprit tortoises show
great pugnacity toward each other, repeatedly ramming one
another with their spadelike gular projections and trying—often
successfully—to turn each other over. James Juvik has colorfully
described these tortoises as acting like animated crowbars.
462
The natural diet of the bowsprit tortoise is unknown. Captive

specimens are normally vegetarian but will sometimes eat meat.
Nesting is known to take place in August; usually a single egg is
laid, but sometimes two. The nest hole is reported to be three or
four inches deep, and the eggs range in size from 1.38 x 1.45 inches
to 1.34 x 1.65 inches. At least in the more southern parts of the
range, incubation may take as long as a year or 14 months.
This species is reported to do quite well in captivity, but al
though common in its habitat it is not often exported. The species
was imported into Britain in large numbers after the Second World
War, but before long the British pet trade switched to southern
Europe and North Africa for its tortoise supplies.
GENUS MALACOCHERSUS
The African pancake tortoise, Malacochersus tomieri, is
surely the most extraordinary tortoise in the world. It appears to
make a point of being what any other tortoise is not; other tortoises
have hard, domed shells and retract head and limbs when dis
turbed. Malacochersus has an extremely flat, sometimes even
swaybacked, shell which is so soft that it can be squeezed between
thumb and finger; moreover the animal, when molested, runs
away and hides itself in a convenient rocky crevice, rather than
retiring into its shell.
The pancake tortoise is found in isolated locations in Kenya
and Tanzania. The southernmost record is from Lindi, on the coast
of Tanzania. The northernmost record is from Mathews Ridge,
Kenya, and the westernmost from just south of Lake Victoria. The
preferred habitat is reported to be areas of small hills (“kopjes”)
with rocky outcrops in arid thombrush, at altitudes from about 100
to 4000 feet (Loveridge and Williams 1957: Bull. Mus. Comp. Zool.,
vol. 115, no. 6). Eglis (1967: Int. Turt. Tort. Soc. J., no. 1)
increased the maximum reported altitude to 6000 feet. The tor
toises appear not to venture far from the particular rocky outcrops
that constitute their “home,” so the various populations are all
reproductively isolated and show considerable variation in pat
tern. Despite the discontinuous range, they may locally be very
abundant. Loveridge found eleven under one slab of rock at
Dodoma.
The carapace of a fully adult pancake tortoise measures about
six inches; the record shell length is seven inches. Females get
slightly larger than males. The scutes of the carapace show no
obviously unusual features except that, because of the flatness of
the shell, the three middle vertebral scutes are relatively small.
There are twelve pairs of marginal scutes and two supracaudals.
The coloration of the carapace is very variable; the typical pattern
involves an overall pale brown color with thick very dark brown or
black scute borders (except on the posterior parts of the scutes,
463
www.Ebook777.com
where very little growth takes place). The dark scute borders are
penetrated by asymmetrical radiating light streaks, the density of
which is very variable.
The nuchal, pygal, and peripheral bones are thin but more or
less normal. However, the flexible middle part of the carapace has
a most unusual structure. The pleural bones have wide spaces be
tween them, such as are present in embryos and hatchlings of
other tortoises. But in contrast to such cases, the ribs disappear
entirely in adult pancake tortoises. The pleural bones are thin and
alternate in width, the third, fifth, and seventh members being
very narrow. The broader second, fourth, and sixth pleural bones
completely fail to make any contact with the neural series, the
neural bones corresponding to these pleurals being extremely nar
row and lacking lateral processes. The adult tortoise thus not only
has a complete series of lateral fontanelles, but also has a series of
four butterfly-shaped fontanelles along the midline. However,
toward the back several neural and suprapygal bones form a con
tinuous row. Despite the excessively fenestrated bony shell struc
ture, at every point the borders between scutes have bone beneath
them.
The plastron of Malacochersus shows wide, truncated gulars
and a strong anterior epiplastral expansion. In certain populations
an intergular scute may be present. The scutes, like those of the
carapace, are pale brown with wide dark brown borders on the
front and sides, penetrated in some places by light radiating mark
ings. The bony structure of the plastron shows no recessed gular
lip, but otherwise is more or less normal at the front and back; but
in the middle an enormous fontanelle is present, occupying per
haps one third of the entire plastral area. This fontanelle is bor
dered at the sides by ventral expansions of the fifth and sixth
peripheral bones, or (at least in one specimen in my possession) by
diamond-shaped laterally-situated mesoplastra and by a down
ward process of the sixth peripheral.
The head of the pancake tortoise is relatively normal, though
the jaw edges are markedly serrated. The skull is well ossified.
The limbs are slender and are unusual in the extreme flexibility,
especially at the wrists.
The various morphological specializations of Malacochersus
serve it well in its peculiar mode of life. When disturbed, it takes
refuge either beneath heavy boulders or in vertical cracks in huge
rocks. When in such a position, it inflates its lungs and thus wedges
itself almost immoveably. Frequently it falls while carrying out its
normal program of rock climbing, and presumably at least half
the time it lands on its back. However, here its flat shell and slen
der, flexible limbs stand it in good stead, and it rights itself without
difficulty.
464
The unique African pancake tortoise, Malacochersus tornieri, is one of the

few turtles that can be found under rocks. Photo by G. Marcuse.
In nature the pancake tortoise is known to eat dry grass. In

captivity it will accept any of a wide variety of fruits and vege
tables.
At the present time only one species of pancake tortoise is
recognized. However, the still unique type specimen of 1Malaco
chersus procterae, strikingly different in shell pattern, body
depth, and head scalation, will eventually require either confirma
tion or formal disproof. A third described species, M. loveridgii,
appears to be simply a local population variant of tornieri. The
relationship of Malacochersus to other tortoises is obscure; Love-
ridge and Williams suggest that it may be independently derived
from the ancestral stock of the testudinines.
The pancake tortoise lays a single egg. In the wild, nesting is
said to take place in July and August, but captive specimens in the
San Diego Zoo have laid in all months of the year. One female may
lay twice or more in a year. The egg is elongate, measuring on
average 1.84” x 1.14”. Twenty-eight eggs in the San Diego Zoo
hatched after an average of 186 days. The hatchlings have a mean
carapace length of 1.55”; they are not quite as flat as the adults
and have more rigid carapaces, but the plastra are quite flexible.
James Juvik has recently demonstrated that pancake tor
toises are capable of climbing up vertical rocky crevices using a
465
www.Ebook777.com
technique that mountaineers call “chimney climbing;” the cara
pace is pressed against one wall of the crack and the feet are
braced against the other. By a combination of wriggling and a
selective application and withdrawal of pressure, the tortoise can
slowly move upward.
Courtship in the pancake tortoise involves the male snapping
at the limbs of the female, presumably to immobilize her before
mounting. Two males described by Loveridge and Williams were
“so ill-tempered as to seise the edges of the females’
carapaces in their jaws, drag them along and eventually get
beneath as they apparently attempted to overturn them. ”
These authors mention the mating season as being in January
and February.
GENUS PYXIS
The little Malagasy spider tortoise, Pyxis arachnoides, is the
only tortoise with the anterior lobe of the plastron hinged. This
attractive species has a rather domed carapace, black in color,
with a yellow margin on each of the vertebral and costal scutes as
well as a pattern of radiating yellow lines. These light rays are
fewer in number than is normal among “starred” tortoises, and on
the costals they tend to be concentrated about a longitudinal axis,
being absent from the areas directly above and below the areolae.
The general effect, reminiscent of a spider’s web, gives this spe
cies its common and scientific names. The plastron is unpigmen-
ted except for a few dark spots on the bridge that are not invari
ably present. The head is predominantly black, with small light
areas. The limbs are yellow on the outer surfaces and black on the
inner ones.
The plastral hinge of Pyxis is located between the epiplastral
and hypoplastral bones and between the humeral and pectoral
scutes. The hinge passes across the rear of the entoplastron, which
thus lacks the posterior spine present in most turtles. The margin
of the moveable lobe is not rounded as in the box turtles, but has a
distinctly wavy edge, with a noticeable emargination between the
gulars. The closure is thus not as thorough as in a box turtle.
It appears that the cervical column of Pyxis is thrown into an
unusual conformation when the head is retracted and the anterior
plastral lobe raised, and this is reflected in the cervical vertebrae.
Pyxis is the only tortoise in which the cervical vertebrae (except
for the atlas) may be procoelous throughout the series, i.e. have an
anterior articular concavity and a posterior convexity in all cases.
This condition was found in 7 out of 88 specimens examined by
Ernest Williams. The one other known cervical pattern for this
species, found in the remaining 81 specimens, includes a biconvex
466
second centrum. Such a pattern is only known as a rare variant in

other turtles.
What appears to be an abnormal specimen of Pyxis in which
the plastral hinge completely failed to develop was recently made
the type specimen of Pyxoides brygooi by Simone Vuillemin and
Charles Domergue.
Pyxis is known to occur in the extreme southern part of Mada
gascar, where it is locally called “kapila.” It has also been found
at Tulear and at Majunga, and toward the north-east as far as the
mouth of the Mahajamba River. Little is known of its habits, and it
is difficult to separate fact from nonsense in the available writings
about this species. It is often called the smallest of all tortoises, but
in fact it can reach a carapace length of over six inches—distinctly
bigger, therefore, than several South African tortoises of the
genera Homopus and Psammobates. It is also often stated to be
among the rarest of the tortoises; but if this is so, it is difficult to
see how Voeltzkow managed to collect no fewer than 630 speci
mens on his expedition to Madagascar in the late nineteenth cen
tury. This enormous series was deposited in the Vienna Natural
History Museum. My correspondent John Visser informs me that
the species is relatively abundant in southwestern Madagascar.
However, it is true that the species is very rarely seen in zoological
gardens or in private collections.
The skull of Pyxis has recently been described by Vuillemin
and Rabodomihamina (Bull. Soc. Zool. France, 93, No. 1., 1968, pp.
25-31), and several major variations from the skulls of other tor
toises were claimed. However, from the figures it is clear that the
“pseudo-temporal fenestra” described is simply the normal skull
roof emargination present in all tortoises, and the supposed supra-
temporal bones are simply the pro-otics, again present in all tur
tles. Divisions were present in the skull figured in the prefrontal
and postfrontal bones, but it is very unlikely that the fragments
split off are truly homologous with the nasal and postorbital bones
of primitive reptiles, as the authors claim.
Structurally, Pyxis shows the alternating octagonal-square
neural bone pattern typical of advanced tortoises. However, al
though this would seem to suggest a relationship to Geochelone,
Testudo, Gopherus, etc., it is more probably an independently
evolved modification, the true affinities of this genus lying with
Acinixys, Kinixys, Homopus, etc. The pygal pattern of Pyxis is
primitive, i.e. does not involve one suprapygal bone enclosing a
smaller one between its ventral rami. The jaw surfaces of Pyxis
show feeble denticulations but lack a maxillary triturating ridge.
The failure of the quadrate to enclose the stapes in Pyxis is an
emydine character ; among other tortoises it is only found in Tes
tudo kleinmanni and Geochelone sumeirei. In Pyxis, the costomar-
467
www.Ebook777.com
ginal sulcus tends to be located well down on the peripheral bones,
as in Kinixys and Acinixys, but in contrast with other tortoises, in
which the costomarginal sulcus is more or less congruent with the
pleuro-peripheral suture.
According to Siebenrock (1910: Sitzungsbr. Akad. Wiss. Wien,
119, no. 1, pp. 693-720), not only is the anterior plastral lobe of Pyxis
fully flexible, but in addition the posterior lobe shows some spon
taneous mobility during oviposition.
GENUS ACINIXYS
Acinixys ylanicauda, like Pyxis, is confined to the island of
Madagascar. It is a small species, approximately five inches long
when fully grown. It has a somewhat lower and more elongate
carapace than Pyxis and lacks the anterior plastral hinge. The lar
ger scutes of the carapace have broad yellow margins and yellow
or light brown areolae. Surrounding the areola of each scute is a
thick black band, corresponding in shape to the overall shape of
the scute. The black band on each scute is penetrated by a small
but variable number of light rays. As in Pyxis, in many specimens
the light rays on the costals are clustered around the longitudinal
axis. The marginal scutes are decorated with light vertical
streaks. The pattern is well defined and highly contrasting in
young specimens, but becomes less distinct in old ones. A short,
broad nuchal scute is usually, but not invariably, present. The ver
tebral region is typically flattened, but a specimen from Moron-
dava, on the south-west coast of Madagascar, with slightly bulbous
vertebral scutes, has recently been declared the type specimen of
a new species, “Testudo morondavaensis’’ Vuillemin 1972. The
carapace margin of Acinixys is somewhat serrated in young ani
mals, particularly anteriorly. The shell outline, when viewed from
above, is either almost perfectly oval or slightly narrowed anteri
orly.
The plastron of Acinixys is unpigmented except for a few dark
blotches situated towards the sides, particularly around the ab
dominal areolae. The gulars are separated anteriorly by an angu
lar notch. The posterior shell opening is very narrow, but deposi
tion of the rather large eggs is made possible by a slight mobility
between the abdominal and femoral scutes in adult females (as in
some large female Testudo), as well as some flexibility along the
interfemoral and interanal seam. Plastral mobility is thus situated
in front of and between the xiphiplastra.
The head is mainly dark brown to black, but there are some
light spots, and the upper mandibular plate is entirely unpigmen
ted. The limbs are covered with yellow scales separated by skin.
The tail is flattened and has a large fingernail-like scale on the
end. The vent of the male opens almost at the tip of the tail.
468
Structurally, Acinixys is primitive in having hexagonal neural

bones and also in retaining a maxillary ridge (lost in Pyxis, Kin-
ixys, etc.). The pygal pattern is primitive: there is no large supra-
pygal partially enclosing a smaller one between its ventral rami.
Also, the pleuroperipheral suture is situated well above the costo-
marginal sulcus—another primitive character.
The habits of Acinixys are virtually unknown. Even the range
is imperfectly defined, but most references give it as western or
southwestern Madagascar. The type locality of both Acinixys
planicauda and of its synonym Testudo morondavaensis is given
as Morondava, or Mouroundava. The egg of Acinixys is relatively
large and only slightly ovoid; it weighs between 15 and 20 gm.
The natural abundance of Acinixys is unknown, but the species
was for a long time unknown in the living state in the New World.
However, the San Diego Zoo received six living specimens in 1970,
and James Juvik of Hilo College, Hawaii, has recently obtained
some specimens for captive breeding purposes.
BACKGROUND READING
Au ffenberg, W. 1974. Checklist of fossil land tortoises (Testudin-
idae). Bull. Florida State Mus., 18(3): 121-251.
Eglis, A. 1967. Tortoise behaviour: a taxonomic adjunct. Int. Turt.
Tort. Soc. J. 1(2): 6-9, 33.
Loveridge, A. and E.E. Williams. 1957. Revision of the African tor
toises and turtles of the suborder Cryptodira. Bull. Mus. Comp.
Zool, 115(6): 163-557.
Williams, E.E. 1950. Testudo cubensis and the evolution of Western
Hemisphere tortoises. Bull. Amer. Mus. Nat. Hist. 95, 1: 1-36.
__________ 1952. A new fossil tortoise from Mona Island, West
Indies, and a tentative arrangement of the tortoises of the
world. Bull. Amer. Mus. Nat. Hist. 99(9): 541-560.
GEOCHELONE SULCATA:
Flower, S.S. 1928. Exhibition of a great African tortoise (Tes
tudo sulcata). Proc. Zool. Soc. London, p. 654.
__________. 1937. Further notes on the duration of life in ani
mals. Ill Reptiles. Proc. Zool. Soc. London, pp. 1-39.
Switak, K. 1972. Tortoise keeping at its best. Int. Turt. Tort.
Soc. J. 6(4): 20-25,39.
caise. I.F.A.N., Dakar. Pp. 1-354.
GEOCHELONE PARDALIS:
Archer, W.H. 1968. A classification problem. African Wild Life
22(3): 249-254.
469
www.Ebook777.com
Hewitt, J. 1937. A guide to the vertebrate fauna of the Eastern
Cape Province, South Africa. Part II. Reptiles, Amphibians,
and Freshwater Fishes. Grahamstown, pp. 1-118, App. A-B,
G-H, pis. i-xxviii.
Loveridge, A. and E.E. Williams. 1957. op. cit.
Rowe, J and J. Janulaw. 1971. A noteworthy conservation
achievement. Int. Turt. Tort. Soc. J. 5(4): 20-30.
Switak, K. 1971. In Kruger National Park. Int. Turt. Tort. Soc.
J. 5(1): 6-9,27.
GEOCHELONE ELEGANS and G. PLATYNOTA
Anderson, J. 1878-79. Anatomical and zoological researches and
zoological results of the Yunnan Expeditions. Reptilia and
Amphibia, pp. 705-860. Calcutta.
Beltz, R.E. 1968. The star in your collection. Int. Turt. Tort. Soc.
J. 2(6): 17-19.
Colombo. 412 pp.
Smith, M.A. 1931. Fauna of British India, Vol. 1. London. Pp.
i-xxvi, 1-179.
GEOCHELONE RADIATA and G. YNIPHORA
Auffenberg, W. 1963. Testudo hypselonota Boirret referred to
Geochelone radiata (Shaw). J. Bombay Nat. Hist. Soc. 60
(2): 462-465.
Baudy, R.E. 1970. In quest of Geochelone radiata. Int. Turt.
Tort. Soc. J. 4(1): 19-23, 27.
Juvik, J.O. 1975. The turtles and tortoises of Madagascar. De
fenders of Wildlife, April 1975.
__________ In press. The survival status of Geochelone radiata
in southern Madagascar: a preliminary assessment.
__________ and C.P. Blanc. 1974. The angonoka of Cape Soda.
Animals, 16(4): 148-153.
Paulian, R. 1955. Les Animaux Protegees de Madagascar.
Publ. Inst. Rech. Sci. Tananarive, Madagascar.
Vuillemin, S. 1972. L’articulation cervico-dorsale chez Testudo
(Geochelone) radiata Shaw, 1802 (Chelonien). Ann. Univ.
Madagascar, Serie Sci. Nat. Math. 9: 183-192.
Zovickian, W.H. 1973. Reproduction of the radiated. Int. Turt.
Tort. Soc. J. 7(2): 26-29,35.
GEOCHELONE, subgenera ALDABRACHELYS and
CYLINDRASPIS
Boulenger, G. A. 1893. On remains of an extinct gigantic tortoise
from Madagascar (Testudo grandidieri Vaillant). Trans.
Zool. Soc. London XIII 1894, 305-311.
Carie, P. 1925. Les tortues geantes des iles de l’Ocean Indien.
Comptes Rendus Soc. Biogeographie Paris 13: 91-92.
470
Davidson, W.E. 1911. Land tortoises in the Seychelles. Proc.

Zool. Soc. London 1911: 622.
Deraniyagala, P.E.P. 1960. Two giant tortoises in the Colombo
National Museum. Spol. Zeyl. 29, Pt. 1: 19-22.
Dumeril, A.M.C. and G. Bibron, 1835. Erpetologie generale, ou
histoire naturelle complete des Reptiles. Vol. II, Paris: 1-
673.
Fauvel, A. A. 1900. Textes inedits concernant les tortues de terre
gigantesques de l’ile Juan de Nove (I. Farquar) (Ocean
Indien). Bull. Mus. Paris 6: 170-174.
Feuillee-Bil'ot, A. 1932. Les tortues gigantesques au Museum
National d’Histoire Naturelle. La Nature, Paris 1932, 2882:
492-494.
Frazier, J. 1968. Behavioural-ecological observations on giant
tortoises (Geochelone gigantea) on Aldabra January to June
1968. Unpublished ms.
Fritsch, A. 1871. Zur Anatomie der Elephanten-Schildkrote.
Abhandl. bohm. Ges Wiss. iv 1871: 18 pp.
Froidevaux, H. 1889. Textes historiques inedits ou peu connus
relatifs aux tortues de terre de l’lle Bourbon. Bull. Mus.
Paris 1899, p. 214.
Gadow, H. 1893. On the remains of some gigantic land-tortoises,
and of an extinct lizard, recently discovered in Mauritius.
Trans. Zool. Soc. xiii, 1894: 313-322.
__________. 1901. Amphibia and Reptiles. Cambridge Natural
History, Vol. VIII; i-xiii, 1-668.
Gaymer, R. 1968. The Indian Ocean giant tortoise Testudo
gigantea on Aldabra. J. Zool. London 154: 341-363.
Grimpe, G. 1928. Merkwurdige Todesursache einer Elefanten-
Schildkrote. Zool. Garten Leipzig 1 1928, pp. 225-226.
Grubb, P. 1971. The growth, ecology, and population structure
of giant tortoises on Aldabra. Phil. Trans. Roy. Soc. London
B 260: 327-372.
Gunther, A.C.L.G. 1875a. Preliminary notice of some extinct
tortoises from the islands of Rodriguez and Mauritius. Ann.
& Mag. Nat. Hist. (4) xi: 397.
_________ . 1875b. Description of the living and extinct races of
gigantic land tortoises, pts. i and ii. Phil. Trans, clvi, 251-284.
__________. 1877. The gigantic land-tortoises (living and extinct)
in the collection of the British Museum. London: i-iv, 1-96.
_________ . 1893. The Mauritius gigantic land-tortoise. The
Times, Jan. 12 1893.
__________ 1898. Recent progress in our knowledge of distribu
tion of gigantic land-tortoises, with list of relevant publica
tions, 1875-1898. Proc. Linn. Soc. cx: 14-29.
Haddon, A.C. 1881a. On the extinct land-tortoises of Mauritius
471
www.Ebook777.com
and Rodriguez. Trans. Linn. Soc. (Zool.), 2nd. Ser., II: 155-
164.
__________ 1881b. On the extinct land-tortoises of Mauritius and
Rodriguez. J. Linn. Soc. xv: 58-59.
Hubrecht, A.A.W. 1881. On certain tortoises in the collections of
the Leyden Museum. Notes Leyden Mus. 3: 41-50.
Keynes, Q. 1950. St. Helena: the forgotten island. Nat. Geog.
Mag. 98(2): 265-280.
__________ 1959. Seychelles, tropic isles of Eden. Nat. Geog.
Mag. 116(5): 670-695.
Loveridge, A. 1962. Our tortoises. Part 1. The truth about Jona
than. ‘Wirebird’ 111(88): 704-706.
__________ 1969. St. Helena News Review, 41: Nos. 1557 and
1558.
Lucas, F.A. 1889. The Galapagos and Mascarene tortoises.
Smithsonian Report, 1889: 643-647.
Lydekker, J. 1910. Giant tortoises and their distribution.
Science Progress, 1910: 5, 302-317.
Malavois, M. 1786. Description et notice sur l’Archipel des Sey
chelles. (Manuscript, Vol. 106, in Caen Public Library).
Milne-Edwards, A. 1868. Sur des decouvertes zoologiques faites
recemment a Madagascar par M. Alfred Grandidier.
Compt. Rend. 1868, Pt. 2: 1165-1167.
__________ 1875. Nouveaux documents sur l’epoque de la dis-
parition de la faune ancienne de l’ile Rodriguez. Ann. Sc.
Nat. art. no. 4, pp. 20.
Nigrelli, R.F. 1954. Some longevity records of vertebrates.
Trans. N.Y. Acad. Sci. (2)16 6: 296-299.
Oliver, P. 1891. The voyage of Francois Leguat of Breese to
Rodriguez, Mauritius, Java, and the Cape of Good Hope.
Edited and annotated by Captain P. Oliver. London (Hak
luyt Society), 1891. 2 vols.
__________. 1892. Mauritius and its gigantic tortoise. Illustr.
London News, Dec. 3 1892, p. 715.
Ommanney, F.D. 1952. The shoals of Capricorn. Longmans,
Green and Co., London, i-viii, 1-322.
Parker, H.W. 1928. Two extinct giant tortoises. Nat. Hist. Mag.,
London 1: 152-154.
__________ 1929. A gigantic land tortoise from the Seychelles.
Nat. Hist. Mag., London II, No. 9: 41-43.
Pearson, J. 1910. The Ceylon giant tortoise; the giant tortoise at
Galle. Spol. Zeyl. 7, pp. 109 & 208-209.
Perrault, J. 1676. Description anatomique d’une grande tortue
des Indes. Mem. pour servir a l’Hist. Nat. des Animaux, p.
193.
472
Petit, M. 1737. Description anatomique des yeux de la gren-

ouille et de la tortue. Hist. Acad. Roy. des Sciences, Paris
1741: 142-169.
Quaranta, J.V. 1952. An experimental study of the color vision
of the giant tortoise. Zoologica, N.Y., 37: 295-312.
Rothschild, W. 1894a. On giant land tortoises. Novit. Zool. 1:
676.
__________ 1894b. Remarks and corrections relating to the
living giant tortoise of Mauritius. Novit. Zool. 1: 690.
__________. 1895. Further notes on gigantic land tortoises. Novit.
Zool. 2: 483.
__________ 1896. Further notes on gigantic land tortoises. Novit.
Zool. 3: 85-91.
__________ 1897. Further notes on gigantic land tortoises. Novit.
Zool. 4: 407-408.
__________ 1899. Note on Testudo sumeirei. Novit. Zool. 6: 359-
360.
__________ 1906. A new species of giant tortoise. Novit. Zool.
13: 753-754.
__________ 1915. On the gigantic land tortoises of the Seychelles
and Aldabra-Madagascar group with some notes on certain
forms of the Mascarene group. Novit. Zool. 22: 418-442.
__________. 1925. Giant tortoises of the Galapagos and Masca
rene Islands. Proc. Linn. Soc. London 137: 40.
__________. 1928. Notes on gigantic land tortoises. Proc. Zool.
Soc. London 1928: 658-660.
Sauzier, T. 1892. Tortue de terre gigantesque a l’ile Maurice.
La Nature Nov. 1892: 395-398.
__________. 1893. Les tortues de terre gigantesques de Madagas
car, et de certaines autres iles de la mer des Indes. Paris,
G. Masson. Pp. 1-32.
__________ 1895. Gigantesque tortue terrestre, d’apres un speci
men vivant. La Nature, xxiii: 273-275.
Schneider, J.G. 1783. Naturgeschichte der Schildkroten, pp.
1-364. Leipzig: Johann Gottfried Mullerschen.
Schoepff, J.D. 1792. Historia Testudinum. Erlangae.
Siebenrock, F. 1903. Bemerkungen zu Herm. Dr. P. Schacht’s
Abhandlung: Beitrage zur Kenntnis der auf den Seychellen
lebenden Elephanten-Schildkroten. Zool. Anz. xxvi: 366-368.
Vaillant, L. 1885. Remarques complementaires sur les tortues
gigantesques de Madagascar. Compt. Rend. 1885: 874-877.
__________. 1893. Les tortues eteintes de l’ile Rodriguez, d’apres
les pieces conservees dans les galeries du Museum. Centen.
Mus. d’Hist. Nat. 1893: 255-288.
__________. 1898. Dessins inedits de Cheloniens tires des manu-
scrits de Commerson. Bull. Mus. Paris, 1898: 133-139.
473
www.Ebook777.com
__________ 1899a. Nouveaux documents historiques sur les tor-
tues terrestres des Mascareignes et des Seychelles. Bull.
Mus. Paris 5: 19-23.
__________. 1899b. Documents relatifs a la tortue gigantesque
de la Reunion. Bull. Mus. Paris, 5: 354-356.
__________ 1900. La tortue de Perrault (Testudo indica Schnei
der). Nouv. Archives du Mus. Paris (4ieme Ser.) 2: 25-47.
Voeltzkow, A. 1896. Reisenschildkroten von Aldabra. Zoolog.
Gart. 1896: 30-31.
GEOCHELONE, subgenera MANOURIA and INDOTESTUDO:

Auffenberg, W. 1964a. A first record of breeding colour changes
in a tortoise. J. Bombay Nat. Hist. Soc. 61(1): 191-192.
__________. 1964b. Notes on the courtship of the land tortoise
Geochelone travancorica (Boulenger). J. Bombay Nat. Hist.
Soc. 61(2): 247-253.
Boulenger, G.A. 1912. A vertebrate fauna of the Malay Penin
sula from the Isthmus of Kra to Singapore, including the
adjacent islands. Reptilia and Batrachia, pp. i-xiii, 1-286.
Lekagul, Boonsong. 1966. Tortoises of Thailand. Conservation
News, Special Issue no. 7, pp. 56-59.
Taylor, E.H. 1970. The turtles and crocodiles of Thailand and
adjacent waters. Univ. Kansas Sci. Bull. XLIX(3): 87-179.
GEOCHELONE, SOUTH AMERICA:

Auffenberg, W. 1965. Sex and species discrimination in two
sympatric South American tortoises. Copeia No. 3, 335-342.
__________ 1969. Land of the Chaco tortoise. Int. Turt. Tort. Soc.
J. 3(3): 16-19,36-37.
__________ 1971. A new fossil tortoise, with remarks on the
origin of South American Testudinines. Copeia No. 1,106-117.
Freiberg, M.A. 1971. El mundo de las tortugas. Editorial Alba-
tros, Buenos Aires. 143 pp.
__________ 1973. Dos nuevas tortugas terrestres de Argentina.
Bol. Soc. Biol, de Conception, 46: 81-93.
__________ 1974. The Argentine land tortoise, Geochelone chil-
ensis, an endangered species. Bull. Maryland Herp. Soc.,
10(2): 39-41.
Legler, J.M. 1963. Tortoises (Geochelone carbonaria) in Pan
ama: Distribution and variation. Amer. Midland Nat. 70
(2): 490-503.
Medem, F. 1962. La distribution geographica y ecologia de los
Rev. Acad. Colombiana de Ciencias Exactas, Fis. y Nat. XI,
No. 44: 279-303.
474
Williams, E.E. 1960. Two species of tortoises in Northern South

America. Breviora, Mus. Comp. Zool., No. 120: 1-13.
GEOCHELONE ELEPHANTOPUS:
Baur, G. 1890. The gigantic land tortoises of the Galapagos
Islands. Amer. Nat. 23: 1039-1057.
Beck, R.H. 1903. In the home of the giant tortoise. Rep. N.Y.
Zool. Soc. 7: 1-17.
Beebe, W.L. 1924. Galapagos—World’s End. Putnam’s, New
York.
Carpenter, C.C. 1963. Notes on the behaviour and ecology of the
Galapagos tortoise on Santa Cruz Island. Proc. Okla. Acad.
Sci. 46: 28-32.
Castro, M. 1970. Chatham Island Tortoise. Animals 13(1): IS
IS.
Darwin, C. 1845. Journal of Researches into the Natural History
and Geology of the Countries visited during the Voyage of
H.M.S. Beagle round the world. Second Edition. Ward, Lock
and Co. London.
De Sola, R. 1930. The Liebespiel of Testudo vandenburghi, a
new name for the mid-Albemarle Island Galapagos Tor
toise. Copeia 1930(3): 79-80.
De Vries, T. 1968. On the spot. Int. Turt. Tort. Soc. J. 2(2): 16,
18-19, 33.
Gans, C. and G.M. Hughes. 1967. The mechanism of lung venti
lation in the tortoise Testudo graeca Linne. J. Exp. Biol.
(1967) 47: 1-20.
Garman, S. 1917. The Galapagos tortoises. Mem. Mus. Comp.
Zool. Harv. 30: 261-296.
Gunther, A.C.L.G. 1876. Description of the living and extinct
races of gigantic land-tortoises. Parts I-II. Introduction, and
the tortoises of the Galapagos Islands. Phil. Trans. Roy. Soc.
165: 251-284.
__________. 1877. op. cit.
Hamann, Ole. 1973. Dyr i focus Galapagos oerne—kaempeskild-
passernes national park. Naturens Verden, Copenhagen, 2:
41-51.
Harlan, R. 1827. Description of a land tortoise, from the Gala
pagos Islands, commonly known as the “elephant tortoise.”
J. Acad. Nat. Sci. Philadelphia, 5: 284.
Hendrickson, J.R. 1965. Reptiles of the Galapagos. Pacif. Dis
covery 18(5): 28-36.
__________. 1966. The Galapagos tortoise, Geochelone Fitzinger
1835 (Testudo Linnaeus 1758 in part). In: R.I. Bowman
(Ed.), Proc. Symp. Galapagos Int. Sci. Project: 252-257.
Univ. Calif. Press, Berkeley and Los Angeles.
475
www.Ebook777.com
King, F.W. 1968. Danger—do not move. Int. Turt. Tort. Soc. J.
2(2): 17, 28-30, 35.
MacFarland, C.G. and J. MacFarland. 1972. Goliaths of the
Galapagos. Nat. Geog. Mag., 142(5): 632-649.
------------ and W.G. Reeder. 1974. Cleaning symbiosis involv
ing Galapagos tortoises and two species of Darwin’s finch. Z.
Tierphychol., 34: 464-483.
__________and____________In press. Breeding, raising, and re
stocking of giant tortoises (Geochelone elephantopus) in the
Galapagos Islands. Contr. No. 154 of the Charles Darwin Re
search Foundation for the Galapagos Isles.
__________, J. Villa, and B. Toro. 1974. The Galapagos giant tor
toises (Geochelone elephantopus). Part II: Conservation
methods. Biol. Conservation, 6(3): 198-212.
Perry, R. 1970. Tortoise rearing in the Galapagos Islands.
Zoonooz 42(7): 8-15.
Pritchard, P.C.H. 1971a. Galapagos tortoises, 1971. Herpetol.
Rev. 3(3): 49-51.
------------- 1971b. A further report on Galapagos tortoises. Her
petol. Rev. 3(1): 25-27.
Shaw, C.E. 1961. Another generation of Galapagos giants.
Zoonooz, San Diego. XXXIV(12): 10-15.
------------- 1963. Growth of the Galapagos tortoise. Zoonooz,
San Diego. XXXI(12): 15.
------------- 1967. Breeding the Galapagos tortoise—success
story. Oryx 9: 119-126.
Slevin, J.R. 1959. The Galapagos Islands. A history of their ex
ploration. Occ. Pap. Calif. Acad. Sci. 25, i-x, 1-150.
Snow, D.W. 1964. The giant tortoises of the Galapagos Islands.
Their present status and future chances. Oryx 7: 277-290.
Staedeli, J.H. 1958. New hope for giants of the past. Zoonooz,
San Diego. XXXI(12): 3-4.
__________ 1973. Concern for the Galapagos Tortoises. Int. Turt.
Tort. Soc. J. 7(1): 16-22.
Throp, J.L. 1969. Notes on breeding the Galapagos tortoise Tes-
tudo elephantopus at Honolulu Zoo. Int. Zoo Yearbook 9:
30-31.
Townsend, C.H. 1925. The Galapagos tortoises in their relation
to the whaling industry. A study of old logbooks. Zoologica
4: 55-135.
__________ 1928. The Galapagos Islands revisited. Bull. N.Y.
Zool. Soc., 31(5): 148-169.
__________ 1931. Giant tortoises nearing extinction are being
propagated in the U.S. Sci. Amer., 144: 42-44.
__________ 1937. Growth of Galapagos tortoises, Testudo vicina,
from 1928 to 1937. Zoologica, 22: 289-292.
476
Van Denburgh, J. 1907. Preliminary descriptions of four new

races of gigantic land tortoises from the Galapagos Islands.
Proc. Calif. Acad. Sci. ser. 4, 1: 1-6.
__________. 1914. The gigantic land tortoises of the Galapagos
Islands. Proc. Calif. Acad. Sci., ser. 4, 2: 203-374.
TESTUDO:
Buskirk, J.R. 1967. Turtles of the Holy Land. Int. Turt. Tort.
Soc. J. 1(2): 20-23.
Gadow, H. 1901. op. cit.
Heron, K. 1968. Tortoises in a French garden. Int. Turt. Tort.
Soc. J. 2(1): 18-19,30-33,39-40.
Lambert, M.R.K. 1970. Tortoise drain in Morocco. Int. Turt.
Tort. Soc. J. 4(5): 16-19,35-37.
Mertens, R. 1946. Uber einige mediterrane Schildkroten-Ras-
sen. Senckenbergiana, Frankfurt a. M. 27, 111-118.
Minton, S. 1966. A contribution to the herpetology of West Pakis
tan. Bull. Amer. Mus. Nat. Hist. 134(2): 27-184.
Obst, F. J. and W. Meusel. 1963. Die Landschildkroten Europas.
A. Ziemsen Verlag, Wittenberg Ltherstadt. Pp. 1-52.
Poljakow, A.V. 1946. On the biology of Testudo horsfieldi.
Zborn. trud. Pedag. Insti. Buchara, 33-42. Buchara.
Pritchard, P.C.H. 1966. Notes on Persian turtles. Brit. J. Herpe
tology 3(11): 271-275.
Wermuth, H. 1952. Testudo hermanni robertmertensi n. subst.
und ihr Vorkommen in Spanien. Senckenbergiana 33, 157-
164.
__________. 1958. Status und Nomenklatur der Maurischen
Landschildkrote, Testudo graeca, in SW-Asien und NO-
Afrika. Senckenb. biol. 39, 149-153.
GOPHERUS:
Auffenberg, W. 1966. On the courtship of Gopherus polyphemus.
Herpetologica, 22(2): 113-117.
__________. 1969. Tortoise behavior and survival. Biol. Sci. Cur
riculum Study, Rand McNally, Chicago. Pp. 1-38.
__________and W.G. Weaver, Jr. 1969. Gopherus berlandieri in
southeastern Texas. Bull. Florida State Mus. 13(3): 141-203.
Brame, A.H. and D.J. Peerson. 1969. Tortoise ID. Int. Turt.
Tort. Soc. J. 3(5): 8-12.
Carr, A.F. 1952. Handbook of Turtles. Comstock Publ. Assoc.,
Ithaca. 542 pp.
Grant, C. 1960. Differentiation of the southwestern gopher tor
toises (genus Gopherus), with notes on their habits. Trans.
San Diego Soc. Nat. Hist., 12: 441-448.
477
www.Ebook777.com
Hanson, K.L. 1963. The burrow of the gopher tortoise. Quart. J.
Florida Acad. Sci., 26: 353-360.
Legler, J.M. 1959. A new tortoise, genus Gopherus, from North-
Central Mexico. Univ. Kansas Publ., Mus. Nat. Hist. 11(5):
335-343.
------------ and R.C. Webb. 1961. Remarks on a collection of Bol-
son tortoises, Gopherus flavomarginatus. Herpetologica. 17:
26-37.
Miller, L. 1932. Notes on the desert tortoise (Testudo agassizi).
Trans. San Diego Soc. Nat. Hist., 7: 187-208.
Patterson, R. 1972. Age, Please? Int. Turt. Tort. Soc. J. 6(5):
6-7.
Pawley, R. 1968. The hidden tortoise of Torreon. Int. Turt. Tort.
Soc. J. 2(6): 20-23,36.
Poorman, F. and R. Poorman. 1971. Agassizi vs. berlandieri.
Int. Turt. Tort. Soc. J. 5(1): 14-16.
Woodbury, A.M. and R. Hardy. 1948. Studies on the desert tor
toise, Gopherus agassizi. Ecol. Monogr., 18: 145-200.
PSAMMOBATES:
Archer, W.H. 1967. The geometric tortoise. African Wild Life 21
(4): 321-329.
Auffenberg, W. 1963. A note on the drinking habits of some land
tortoises. Animal Behaviour 11(1): 72-73.
Hewitt, J. 1933. On the Cape species and subspecies of the genus
Chersinella Gray. Part I. Ann. Natal Mus., 7: 255-293.
------------- 1934. On the Cape species and subspecies of the
genus Chersinella Gray. Part II. Ann. Natal Mus., 7: 303-
349.
__________ 1937. A note on the relationship of the Cape genera of
land-tortoises. Afr. Jour. Sci., 33: 788-796.
Juvik, J.0.1971. The status of Psammobates geometricus in the
Western Cape. Int. Turt. Tort. Soc. J. 5(1): 10-13.
__________ 1972. A home for the So. African geometric. Int.
Turt. Tort. Soc. J. 6(3): 26-27.
Lynch, K.B. The beauty of it. Int. Turt. Tort. Soc. J. 2(2): 25, 34.
Rau, R. 1969. Uber die geometrische Landschildkrote (Testudo
geometrica). Salamandra, 5(1/2): 36-45.
__________ 1971. Cape reserve for one of the world’s rarest tor
toises. African Wild Life, 25(3): 95-96.
Turner, E.H. 1971. Psammobates the delicate one. Int. Turt.
Tort. Soc. J. 5(2): 4-5.
HOMOPUS:
Archer, W.H. 1968. The padlopers. African Wild Life, 22(1): 29-
35.
478
Hewitt, J. 1931. Descriptions of some South African tortoises.

Ann. Natal Mus., 6, pp. 461-506.
KINIXYS:
Angel, F. 1941. Sur la presence a Madagascar des tortues du
genre Kinixys. Bull. Mus. Nat. Hist. (Paris), (2)13: 151-152.
Archer, W.H. 1968. The tortoise with a difference. Int. Turt.
Tort. Soc. J. 2(4): 11-13,35-36.
Laurent, R.F. 1956. Contribution a l’herpetologie de la region
des Grands Lacs de l’Afrique Centrale. I. Generalites. II.
Cheloniens. III. Ophidiens. Ann. Mus. Congo, Zool., 48:
1-390.
__________. 1962. On the races of Kinixys belliana Gray. Bre-
viora, Mus. Comp. Zool., No. 176: 1-6.
Schmidt, K.P. 1919. Contribution to the herpetology of the Bel
gian Congo based on the collection of the American Congo
Expedition, 1909-1915. Bull. Am. Mus. Nat. Hist., XXXIX
(II): 385-624.
Siebenrock, F. 1907. Uber zwei Schildkroten aus Kamerun. Ann.
Naturhist. Hofmus. (Wien), 22: 1-8.
caise. I.F.A.N., Dakar. Pp. 1-354.
Vuillemin, S. 1972. Note sur Madakinixys domerguei n. gen. n.
sp. (Testudinidae). Ann. Univ. Madagascar, Ser. Sci. Nat.
Math. 9: 169-181.
CHERSINE:
Archer, W.H. 1967. The angulated tortoise. African Wild Life, 21
(2): 137-143.
MALACOCHERSUS:
Boulenger, G.A. 1920. Une tortue extraordinaire: Testudo love-
ridgei n. sp. C.R. Acad. Sci. Paris, 170: 263-266.
Eglis, A. 1967. The pancake tortoise. Int. Turt. Tort. Soc. J. 1
(2): 29,44-45.
Juvik, J.0.1971. Chimney climber. Int. Turt. Tort. Soc. J. 5(5):
23-24.
Loveridge, A. 1923. Notes on East African tortoises collected
1921-1923, with the description of a new species of
soft-shelled tortoise. Proc. Zool. Soc. London, pp. 923-933.
__________and E.E. Williams. 1957. op. cit.
Procter, J.B. 1922. A study of the remarkable tortoise Testudo
loveridgei Blgr., and the morphology of the chelonian cara
pace. Proc. Zool. Soc. London XXXIV: 483-526.
Shaw, C.E. 1970. The hardy (and prolific) soft-shelled tortoises.
Int. Turt. Tort. Soc. J. 4(1): 6-9, 30-31.
479
www.Ebook777.com
PYXIS:
Angel, F. 1941. Sur quelques variations montrees par le Pyxis
arachnoides Bell. Testudinidae de Madagascar. Bull. Mus.
Nat. Hist. Nat., Paris (2)17: 223-225.
Bell, T. 1827. On two new genera of land tortoises. Trans. Lin-
naean Soc. of London, 15: 395-397.
Malzy, P. 1964. Sur Pyxis arachnoides Bell, tortue terrestre
du Sud de Madagascar. Bull. Mus. Nat. Hist. Nat. Paris.,
36: 441-443.
Siebenrock, F. 1906. Schildkroten von Ostafrika und Madagas
car. In: Voeltzkow, A.: Reise in Ostafrika in den Jahren
1903-1905. Stuttgart, 2: 6-17.
Vuillemin, S. and C. Domergue. 1972. Contribution a l’etude de
la faune de Madagascar: description de Pyxoides brygooi
n. gen. n. sp. (Testudinidae). Ann. Univ. Madagascar, Ser.
Sci. Nat. Math. 9: 193-200.
__________and L. Rabodomihamina. 1968. Originalite du sque-
lette de “Pyxis arachnoides” Bell, 1827 (Testudinidae). Bull.
Soc. Zool. France, 93(1): 25-31.
Williams, E.E. 1950. Variation and selection in the cervical cen
tral articulations of living turtles. Bull. Amer. Mus. Nat.
Hist., 94(9): 505-562.
ACINIXYS:
Grandidier, G. 1867. Liste des Reptiles nouveaux decouverts en
1866 sur le Cote sud-ouest de Madagascar. Rev. et Mag. de
Zool., p. 233.
Vuillemin, S. 1972. Note sur Testudo morondavaensis n. sp.
Ann. Univ. Madagascar, Ser. Sci. Nat. Math. 9: 127-134.
480
Chapter 6
MUD, MUSK, AND SNAPPING TURTLES
In this chapter we shall consider a number of exclusively New
World turtle species of generally drab coloration and usually ag
gressive and predatory disposition. The six genera included in this
group are: Chelydra and Macroclemys (the snapping turtles);
Kinostemon (the mud turtles); Stemotherus (the musk turtles);
Staurotypus (the giant Middle American musk turtles); and
Claudius (the narrow-bridged mud turtle). Some authors place all
of these genera in a single family, the Chelydridae, but this classi
fication obscures the considerable and fundamental structural
diversity, particularly in the plastron, which these turtles show,
and in fact it is probable that the snapping turtles are not nearly as
closely related to the mud and musk turtles as is often supposed.
The classification we shall use here is to include the two snapping
turtle genera in the family Chelydridae and the four mud and
musk turtle genera in the Kinostemidae. Kinostemon and Ster-
notherus are clearly closely related and in fact barely separable
genera; however, Staurotypus and Claudius are so distinctive that
one could present a cogent argument for the inclusion of each in a
separate subfamily, at least.
Only two full species of snapping turtle are recognized, Chely
dra serpentina, the common snapper, and Macroclemys tern-
mincki, the alligator snapper. These are the preferred vernacular
names, although the “loggerhead” or “loggerhead snapper” is
used for both species in some areas. This name is best repressed,
since not only does it fail to differentiate between the two kinds of
snapping turtle, but it also raises the possibility of confusion with
the loggerhead sea turtle, Caretta caretta, and with the logger-
head musk turtle, Stemotherus minor.
GENUS CHELYDRA
The common snapping turtle, Chelydra serpentina, is a very
widespread and often very abundant species whose range covers
the entire eastern United States, the western margin of its range
extending from central Montana southward to eastern New
Mexico. There are introduced populations in California. In Canada
the snapping turtle is found from Nova Scotia, New Brunswick,
and southern Quebec westward across southern Ontario (where it
481
www.Ebook777.com
Amazonian Indian with arrau turtle that he captured for sale to turtle mer
chants. Photo by Harald Schultz.
482
A Brazilian Indian excavating the eggs of a turtle that has made its nest in the
sandy banks of the Amazon River. Photo by Harald Schultz.
Adult female specimens of Podocnemis expansa captured in the Amazon

River for eating. Photo by Harald Schultz.
483
www.Ebook777.com
extends northward to the Lake Nipissing area), and across ex
treme southern Manitoba to southeastern Saskatchewan. The
exact southern limit of the range in southern Texas is obscure but,
though the species definitely enters northern Coahuila in Mexico, I
do not know of a single record from the coastal State of Tamau-
lipas. In tropical Mexico, however, snapping turtles pick up again
in Veracruz and are caught and sold for food in large numbers in
the markets in Veracruz City and Alvarado. Snapping turtles have
also been recorded from northern Chiapas and have been found in
the Caribbean drainages of the Central American countries,
though I know of only one record from Belize, and the species
appears to be rather scarce throughout, except perhaps locally in
Costa Rica. In Panama the species invades the Pacific drainage,
and snappers have been found in Colombia west of the Andes and
occur quite abundantly in western Ecuador, where large numbers
are sold for food in the Guayaquil market.
The common snapping turtle frequently exceeds one foot in
carapace length. The record length is 18.5”. A very large wild-
caught specimen from Florida that I measured some years ago
had a carapace 17’ ’ in length and 14.5” in width. The weight was 42
lbs. In captivity, specimens of even less shell length than this can
be fattened up to considerably greater weights, and there is one
record of an artificially fattened individual in New England reach
ing 86 lbs. Pope (1939) figures a 62 lb. snapper from Plymouth
County, Massachusetts; this may be a record for a wild-caught
specimen. The maximum size reached by this species does not
The common
snapping turtle,
Chelydra serpen
tina.
484
appear to show geographic variation; unusually big specimens

have been found in Canada, in Ecuador, and many places in be
tween.
The common snapping turtle has a rather broad and flat cara
pace whose sides may be more or less parallel or somewhat diver
gent posteriorly. The front and side margins are smooth, but the
rear margin is coarsely serrated throughout life. The shell of the
young is very rough, with radial rugosities and three tuberculate
keels. However, the shell becomes smooth with maturity, though
an areolar tubercle may persist on each of the vertebrals and cos-
tals. The nuchal scute is much wider than long; there are 22 mar
ginal scutes and a pair of supracaudals. The seam between the
supracaudals may be so short that the fifth vertebral almost
reaches the margin of the carapace. The plastron is highly re
duced, the bridges being unusually long and narrow. The inguinal
scute is in broad contact with the axillary on each side. The
greater part of each bridge is covered with what is presumably an
abdominal scute; there are only five pairs of plastral scutes which
have median lines of contact, so that it may be assumed that the
scutes which partially interpose themselves between the third and
fourth pairs of plastral scutes are displaced abdominals.
The head of the common snapping turtle is large, very flat,
and roughly triangular in shape when viewed from above. The
mouth is very large, with jaw surfaces adapted for simple cutting
rather than for crushing or grinding, and there is a pair of barbels
on the chin, sometimes with smaller ones behind. The neck is very
long and highly papillose. The bases of the four limbs are massive
ly muscular, and the front surfaces of the forelimbs are studded
with large, curved scales. The feet are webbed and the claws are
long and thick. There are numerous soft papillae in the axillary
region and on both fore and hind limbs. The tail is extremely long,
often equalling the shell length in very young specimens, and is
armored dorsally with three rows of tubercles.
The carapace of the common snapping turtle is brown in color,
and the dorsal surfaces of the limbs are a drab grey-brown. The
plastron is basically yellow, while the soft skin underneath may be
light or dark in color—on occasions even reddish. There is often a
vague light area behind the eye, and the jaws are marked with ver
tical dark streaks.
Internally, Chelydra is characterized by a relatively thin cara
pace with an uninterrupted series of neural bones which are typi
cally wider than long. The neurals are often irregular in shape; the
second is frequently octagonal. The nuchal bone has long, curved
processes extending from its anterior margins to reach as far back
as the third peripherals. The proximal ends of the ribs curve
sharply away from the internal surface of the carapace, forming a
485
www.Ebook777.com
White-lipped mud turtle, Kinosternon leucostomum.
Kinosternon scorpioides from the Azuero Peninsula in Panama.
Kinosternon hirtipes. Photo by Dr. C. M. Bogert.

Plastral view of Kinosternon angustipons from Tortuguero, Costa Rica. This

rare species is characterized by the small plastron and bulbous snout; it was
first described in 1965.
Red-spotted mud turtle, Kinosternon scorpioides cruentatum, from Central

America. This mud turtle can close its shell as tightly as a box turtle can.
487
www.Ebook777.com
longitudinal tube in which lie the long and powerful longissimus
dorsi muscles which give the turtle its ability to extend its neck
and head very rapidly. On each side the line along which the ribs
swing away from the shell is marked externally by a shallow longi
tudinal groove which lies along the line which separates the verte
bral scutes from the costals. The plastron has the usual comple
ment of nine bones, but it is connected with the sides of the cara
pace by cartilage only. The entoplastron is arrowhead-shaped, and
there are median fontanelles before, between, and behind the hyo-
plastra and hypoplastra. In the neck, the second and third cervi-
cals are opisthocoelous, the fourth is biconvex, and the remainder
procoelous. The articular surfaces in front of the seventh and
eighth cervicals are usually doubled, but occasionally all joints are
single—a very rare circumstance among cryptodires.
Sexual dimorphism in the snapping turtle is not marked, and
there is no great difference, if any, in adult size between the sexes.
However, the vent of the male is located more posteriorly than that
of the female, and the male also tends to have a smaller plastron
and narrower bridges than the female, presumably to assist in
holding the female during copulation.
When handled, snapping turtles sometimes emit an odiferous
secretion from glands in the inguinal region.
Four subspecies of Chelydra serpentina are currently con
sidered valid: C.s. serpentina, occupying the entire range of the
species in the United States except for the area occupied by C.s.
osceola, i.e. peninsular Florida and the Apalachicola River system
in southern Georgia; C.s. rossignoni, from central Veracruz,
Mexico, south through Guatemala and southern Belize to Carib
bean drainages of Honduras; and C.s. acutirostris, from northern
Honduras through eastern Nicaragua and Costa Rica to Panama,
and lowland Pacific Colombia and Ecuador, south to the Gulf of
Guayaquil. C.s. serpentina and C.s. osceola appear to intergrade
in the Okefenokee Swamp of southern Georgia. These two races
are differentiated by a number of characters; osceola has a deeper
carapace than serpentina, which slopes off fairly sharply at the
rear, and the neck extensor muscles are considerably more devel
oped. The greater muscular development means that the spaces
under the free rib-ends of osceola are much greater than those of
serpentina, and the width across the free ribs is thus wider than
the skull in osceola, narrower than the skull in serpentina. This dif
ference is manifested externally by a lateral displacement in os
ceola of the shallow longitudinal grooves in the carapace men
tioned earlier. Since this groove also marks the boundary between
the central and lateral scutes, the centrals of osceola are wider
than those of serpentina; in quantitative terms, the width of the
488
The pointed papillae on the neck of Chelydra serpentina osceola help

distinguish it from the typical subspecies.
3rd central/height of 2nd costal averages about 0.838 in serpentina,

about 0.973 in osceola.
The Florida snapping turtle also differs obviously from the
northern race in having granular scales on top of the head and
numerous compressed, pointed papillae on the neck and soft
undersurfaces. The northern race has flat scales on top of the
head, fewer in number than those of osceola, and has rounded pim
ples on the neck and soft parts instead of elongate papillae. Some
other differences also separate the two forms, including the more
pointed snout of osceola, numerous pelvic differences, and the
more central placement of the knobs on the fourth and fifth central
scutes of osceola; these are adjacent to the posterior scute mar
gins in serpentina.
Interestingly, the race from southern Mexico and northern
Central America, C.s. rossignoni, shows greater similarity to
osceola than to the geographically closer serpentina. Feuer has
489
www.Ebook777.com
Scorpion mud turtle,
Kinosternon scorpioides
from
scorpioides,
Guyana. The species has
a very wide range.
Sonoran mud turtle, Kinosternon sonoriense. This is the largest mud turtle in
the United States.
490
Eastern mud turtle, Kinosternon subrubrum subrubrum, from DeKalb County,

Georgia.
mawi, a large, primitive, highly aquatic turtle from southern

Dermatemys
Mexico, Guatemala, and Belize. Photographed at the Brookfield Zoo,
Chicago.
www.Ebook777.com
hypothesized that the Mexican and Florida populations were once
continuous around the southern lowlands of the Gulf States, but
that Pleistocene cold conditions in this area forced a southward
migration, and hence division of the stock. The more cold-tolerant
serpentina then invaded the void. If the dermal papillae of the
snapping turtle are devices to increase the surface area of the skin
to facilitate oxygen exchange through the skin, as is the case with
the musk turtles (Stemotherus), then it becomes obvious that the
southern populations (rossignoni and osceola), living in warm
water with less dissolved oxygen, would require a greater develop
ment of the dermal papillae than would the more northern serpen
tina. However, this speculation is complicated not only by the fact
that the whole of the eastern United States is hot in summer, but
also by the fact that the race from equatorial regions, acutirostris,
lacks the well-developed papillae of rossignoni and in fact is mor
phologically very similar to serpentina.
The common snapping turtle is a highly aggressive animal,
well deserving its vernacular name; when picked up or otherwise
molested, the animal’s head is shot forward with incredible speed,
while the mouth closes hard, often with an audible crunch, as the
neck reaches its full extension. However, specimens of all sizes
may be handled quite safely by the long, thick tail, though it is
important to hold the animal away from one’s legs. The “snap”
action is not just defensive; this predatory animal typically stalks
its prey with a slow-motion walking movement, then grabs it with
the same eye-blurring strike.
The snapping turtle is a highly aquatic species, and since it
spends most of its time on the bottom of relatively deep bodies of
water (slow rivers and lakes) it. may be much more abundant in a
given locality than casual inspection of the area would suggest.
However, it may be caught quite easily on hook and line. Occasion
ally individuals will climb out on half-submerged logs to bask, but
much less frequently than most emydid turtles. On land the gait is
slow and deliberate, with the body held high off the ground. This
species has a high tolerance for cold, and in the north specimens
are sometimes seen walking about under ice.
The diet of the snapping turtle in Michigan includes fish, car
rion, invertebrates of various kinds, and a surprising quantity of
vegetable matter. Large snapping turtles will also, on occasion,
seize swimming water fowl by the feet and drag them under for
consumption. In Chiapas, Alvarez del Toro reports that rossignoni
feeds mostly on fish and frogs, but will also eat other small aquatic
animals and any small reptiles and mammals which may come
within its reach. In Colombia, Medem reports that acutirostris
feeds on fish, frogs, crabs, vegetable matter, and molluscs; the
stomach of one male specimen contained the opercula of 101 fresh
water snails.
492
In the United States, snapping turtles copulate from late April

to November. The carapace of the female being quite flat, the
male is able to hold on to the margins of her shell with the claws of
all four feet while the tail is twisted around that of the female until
contact between the vents is established. Nesting, though concen
trated in the month of June, may take place from late April to
October. The excavation of the nest is carried out in the normal
way with alternating actions of the hind feet. The eggs are report
edly directed individually to the bottom of the nest by one of the
hind feet. The average clutch numbers about 25 eggs, spherical or
very slightly ovoid in shape, and a little more than one inch in
diameter. The shells are hard, but tough rather than brittle, so that
the eggs may be dropped several feet onto a hard surface without
breaking. There is some evidence that more eggs are laid in nor
thern parts of the range than in the southern parts, and there are
records of clutches numbering 77 and 80 eggs respectively from
Manitoba. A possible (though unverified) explanation of this is
that southern snappers enjoy a long enough breeding season to be
able to deposit more than one clutch each year, while in the north
eggs have to be laid in early summer if they are to have a good
chance of hatching. On the other hand, some southern snappers
produce quite large clutches. John McPhee (New Yorker, April 28,
1973, p. 45) mentions a snapper about 14” long found injured on the
road east from Hunger and Hardship Creek, Georgia; this turtle
was dissected by Carol Ruckdeschel and found to contain 56
shelled eggs. In Chiapas, the “tortuga lagarto,” as this species is
locally called, lays twenty to thirty eggs, the season lasting from
April to June, while in Colombia, 20 to 35 eggs are laid in February.
In the United States many nests are made at considerable dis
tances from the nearest water, but Medem reports that in Colom
bia nests were between 90 cms. and 3 meters from the nearest
water.
Hatching commonly takes place after 80 to 90 days, but occa
sional overwintering of the young has been observed in the ex
treme northern parts of the range. The hatchlings are slightly over
an inch in length and almost as wide as long. They differ from the
adults in having much rougher shells, dark mottling on the head
and jaws, and frequently white spots on the upper and lower mar
ginals. They defend themselves with strong bites even before they
are completely free of the egg.
GENUS MACROCLEMYS
The only other species included in the family Chelydridae is
the alligator snapping turtle, Macroclemys temmincki. Despite
the superficial similarities between the two forms, generic separa
tion is clearly valid on the basis of several factors. For example,
the carapace of Macroclemys retains three strong ridges through-
493
www.Ebook777.com
Giant musk turtle, Staurotypus triporcatus, from Belize. This is the largest
turtle in the family Kinosternidae. Freshly captured specimens are highly ag
gressive. Photographed at Ross Allen’s Reptile Institute.
Yellow mud turtle, Kinosternon fiavescens, from Texas and Oklahoma. Photo
by Dr. Herbert R. Axelrod.
494
Stink-pot turtle, Sternotherus odoratus, found throughout the eastern

United States.
Southern musk turtle, Sternotherus minor minor, from the Chipola River in
Florida.
495
www.Ebook777.com
out life, and there are between three and five small extra scutes,
known as supramarginals, between the first three lateral scutes
and the true marginals. The snout is more pointed, when viewed
from above, than that of Chelydra, and the eyes are placed on the
sides of the head, rather than toward the top as in Chelydra. The
eye is surrounded by a star-shaped arrangement of fleshy, fila
mentous “eyelashes,” and the plastral scutes often become so
complicated and subdivided that it is wise to give up the task of
trying to name or homologize them. The top of the head is divided
into distinct scales, there being a very large parietal scale sur
rounded by the smaller frontal, supraocular, and temporal scales.
The neck is even more coarsely papillose than that of Chelydra.
The head is extremely large and the neck relatively short, so that
Macroclemys, like Platystemon, has a broad notch in the front of
the carapace to allow the head to be raised for respiratory and
other purposes.
The alligator snapping turtle is much more restricted in its
range than the common snapper. It is found throughout Louisiana
and Arkansas and in adjacent parts of Tennessee, Kentucky, Mis
souri, Kansas, Oklahoma, and Texas. The range covers most of
Mississippi except for the extreme north-east, as well as the
southern half of Alabama, the Okefenokee Swamp area of Georgia,
and the panhandle of Florida, reaching as far as the Suwannee and
Sante Fe Rivers. The species has been recorded in Illinois as far
north as Rock Island County on the Iowa state line.
The alligator snapping turtle, though on occasion a scavenger
or prowling predator like Chelydra, is famous for a much more
subtle method of obtaining food. The animal’s usual habitat is
dark, slow-moving water, where it pursues a way of life so seden
tary that a thick growth of algae forms on the rough carapace. The
outlines of the head and limbs are so irregular and broken up that
the immobile animal becomes practically invisible to passing fish.
The animal holds its huge mouth open astonishingly wide for long
periods of time—even this trait does not give the animal away, as
the interior of the mouth, instead of being pink or white, is greyish
with black blotches. However, on the floor of the mouth there is a
bright pink bifurcated fleshy filament that looks for all the world
like a small worm. The illusion is heightened by the turtle’s ability
to wriggle and move the “worm” most realistically. Fish entering
the trap-like jaws to investigate the bait are either swallowed
whole, sliced in two by the sharp jaws, or are impaled on the sharp,
hooked tips with which both upper and lower jaws are equipped.
That the diet is not exclusively fish, however, is evident from Ross
Allen’s observation that fresh-caught specimens from the Suwan
nee River often void large quantities of clam and mussel shells.
496
Definitely lateral eyes and very strong carapace keels are hallmarks of the
alligator snapping turtle, Macroclemys temmincki. Photo by Dr. Herbert R.
Axelrod.
When handled, the alligator snapper does not make violent

striking movements like the common snapper, but stays remark
ably still with the mouth wide open, periodically everting the
bright white lining of the glottis, though it does stretch round to
bite any part of its handler that is held within reach. Because of the
relatively short neck, an alligator snapper of any size can be held
quite safely with one hand gripping the shell above the head and
the other holding the supracaudal region.
The alligator snapping turtle, according to James Dobie s
recent survey in Louisiana, reaches sexual maturity at a carapace
length of around fourteen inches, when the age (estimated by
counting scute annuli) is 11 to 13 years. At this length the weight of
the animal is between fifteen and twenty pounds. Males reach sex-
497
www.Ebook777.com
Hatchling common snapping turtle, Chelydra serpentina. Photo by Dr.
Herbert R. Axelrod.
Common snapping turtle, Chelydra serpentina. This is one of the largest and
most familiar North American turtles. Photo by Dr. Herbert R. Axelrod.
498
Young alligator snapping turtle, Macroclemys temmincki. The alligator snap

per is the largest freshwater turtle in North America. Photographed at the
London Zoo.
Florida snapping turtle, Chelydra serpentina osceola. This subspecies grows

slightly larger than the common snapper. Photographed at Ross Allen’s Rep
tile Institute.
www.Ebook777.com
ual maturity at a marginally greater size than females. However,
growth continues after maturity is reached, and the large size ob
tained by old specimens places the alligator snapper as the largest
freshwater turtle in the United States—and one of the largest in the
world, especially when one considers that the other really big
freshwater turtles (Dermatemys mawi, Orlitia bomeensis, Podo-
cnemis expansa, etc.) have much smaller heads and extremities
than Macroclemys. The largest female alligator snapper on defi
nite record weighed 51 lbs., but males, which may be distinguished
by their more distally located vents, may get much larger than
this. In 1961, Jess Grammer of Grammer’s Trading Post, Grand
Tower, Illinois, netted a 167 pound Macroclemys upstream from
the Mississippi River near Grand Tower, Jackson County. This
specimen was kept alive and was reported to feed only in the sum
mer months, consuming about 110 pounds of fresh meat annually.
A large specimen from Metropolis, Massac County, Illinois,
weighed 103 lbs. and had a carapace length of 24.02 inches, cara
pace width of 18.19 inches, and head width of 7.87 inches. Philip W.
Smith records a 90-pound individual from Tennessee. There is a
huge specimen alive in the Brookfield Zoo, Chicago, which is sup
posed to weigh 236 lbs. The greatest weight mentioned in the
literature, although one that has been discounted by many
herpetologists, is a 403-pound specimen reported by Hall and Smith
from the Neosho River, Cherokee County, Kansas, in 1937.
In view of the unprecedented size of this specimen, I wrote to
Dr. Smith recently and received the following reply:
“I did not see the Jf03 lb. Macroclemys—the record is
solely dependent upon the memory of Henry Hall, now dead
I understand. He was near retirement when I met him in '45.
Although he was a college biology teacher, I am not confi
dent that all of the bizarre observations he is supposed to
have made are totally unembellished. There is no objective
basis for this feeling, but it inevitably colors my view of his
more extraordinary records. He was a sort of a character—
a raconteur—and not particularly given to meticulous pre
ciseness. I have no doubt that there was a very large alliga
tor snapper, but I fear he may have merely accepted the
exhibitor’s statement of weight—or that his memory failed
him. I have no reason to think that he would deliberately
misrepresent, but I do think he tended to some extent to
accept as truth that which he would like to be true. ”
In view of this doubt, the Brookfield Zoo specimen must be
considered as the largest alligator snapper on definite record.
The largest alligator snapper skeleton I have seen had a skull
width of 9U inches and a carapace length of 31 inches. This was an
500
individual from White Springs, Florida, that was kept alive for a
time at the Everglades Wonder Gardens at Bonita Springs. It
probably weighed over 200 lbs. when alive.
It is very interesting to note that so many of these huge alliga
tor snapper records are from states at the northern extreme of the
animal’s range, where it is extremely rare. For example, there
are only fifteen records available for the whole of Illinois. It is pos
sible that the species does not breed in these northern parts of the
range, and the specimens that occur there have spent their long
lives (very possibly over a hundred years in some cases, since this
is not a particularly rapid-growing species) gradually moving up
river, ending up many hundreds of miles north of their birthplaces.
Indeed, M.M. Wickham actually recorded an Oklahoma specimen
which moves an average of about six miles upstream each year for
three years. A tendency to sit on the bottom of the rivers and to
face upstream (a position that might bring the greatest number of
fish within sight of the “lure” in the mouth) would mean that the
turtle, whenever it walked, would proceed in an upstream direc
tion. As further material to justify this upstream-wandering hypo
thesis, alligator snappers from the Okefenokee Swamp are often
much larger in size than those found downstream in the Suwannee
River, which drains the western side of the swamp; and the ani
mal’s absence from the St. Mary’s River, which drains the eastern
side of the swamp, is inexplicable unless we assume that alligator
snappers of large size are in some way programmed to resist
downstream movement. On the other hand, hatchlings almost cer
tainly tend to move downstream, since they lack the size and
weight to fight the current. After some years they may reach a size
and strength to resist further passive downstream drifting and
start moving back upstream, eventually to breed in those parts of
the river where they themselves were hatched.
Perhaps the most extreme example of this hypothesized up
stream wandering combined with enormous size (and presumably
correspondingly great age) is described by Dr. and Mrs. Sherman
Minton in their book Giant Reptiles. This monstrous animal, ap
parently an alligator snapping turtle, was locally known as the
“Beast of Busco,” and it lived in Fulk’s Lake near Churubusco in
northeastern Indiana. It was seen on several occasions but was
never captured. When spotted on March 1, 1949, the animal was
described as being “as big as a dining room table,” and covered
with moss. It seems possible that this turtle had, over a great
many years, wandered up the Mississippi River and finally
reached northeastern Indiana via the Wabash River. Its weight is
unknown, but an eye-witness estimated that it weighed 500 pounds.
The alligator snapping turtle is reported to lay between 16 and
52 eggs, the average, as with the common snapper, being about 25.
501
www.Ebook777.com
Hatchling southern painted turtle, Chrysemys picta dorsalis. Photo by
Eastern painted turtle, Chrysemys picta picta, one of the most common
American turtles. Photo by Dr. Herbert R. Axelrod.
502
Western painted turtle, Chrysemys picta belli. This is the largest and most at
tractively marked race of painted turtles. Photo by Dr. Herbert R. Axelrod.
Mississippi map turtle, Graptemys kohni. Photo by Dr. Herbert R. Axelrod.
503
www.Ebook777.com
In Louisiana, Dobie concluded that it is unlikely that more than
one clutch is deposited in a season and that some females may lay
on an alternate-year basis. Nesting takes place from April to early
June and may be nocturnal or diurnal. The nest is flask-shaped,
and the eggs are deposited both in the expanded base and in the
narrow neck of the cavity. The eggs are almost spherical, typically
about lW in diameter, but occasionally ovoid eggs may be almost
two inches long. The egg shells are hard and tough, with a consis
tency similar to those of the common snapper. The hatchlings are
about 1.6” or 1.7” in length, with tails about 2.5” long. They are
much darker than the adults, almost black in color, and the shell is
exceedingly rough-surfaced. The “lure” on the tongue is already
present at hatching.
The extent to which this highly aquatic turtle can utilize aqua
tic means of respiration is unknown. Ross Allen and Wilfred Neill
found that captive specimens have to rise for air after fifteen to
twenty minutes submergence. Water is certainly taken in and ex
pelled by the mouth when the turtle is under water, but the
function of this may be olfactory rather than respiratory.
The alligator snapping turtle is used for human consumption
in many parts of its range, especially in Louisiana where the spe
cies probably reaches its maximum abundance.
FAMILY KINOSTERNIDAE
The mud and musk turtles of the family Kinosternidae differ
from the snapping turtles in a number of characteristics. They are
usually small in size, with adult carapace lengths of four to seven
inches (with the exception of the giant musk turtles of the genus
Staurotypus); the tail is always short; the plastron is hinged,
except in Claudius; the entoplastron is absent in Kinosternon and
Stemotherus, and the hyo- and hypoplastra are fused into single
units in Claudius; the abdominal scutes are contiguous (or absent
in Staurotypus and Claudius); the plastron in Kinosternon is rela
tively large and rounded; the neural bone series is interrupted by
certain pairs of pleurals, except in most Staurotypus ; the third
cervical vertebra is biconvex; the caudal vertebrae are procoe-
lous (opisthocoelous in the Chelydridae); the median connection of
the pubes and ischia is bony in Kinosternon and Stemotherus; the
hind margin of the carapace is smooth, never serrated; the foot
plate of the columella is conical (flat in the Chelydridae); the basi-
occipital/opisthotic suture is closed (open in the Chelydridae); the
foramen stapedio-temporale is minute (large in the Chelydridae);
the foramen anteriopalatinum is large (minute in the Chelydri-
504
dae); and the eggs are few in number and strongly ovoid in shape
(numerous and spherical in the Chelydridae).
GENUS KINOSTERNON
Kinosternon is the most polytypic genus of the family, with
probably fourteen full species, several of these with subspecies.
Turtles of this genus are easily recognized by the doubly-hinged
plastron; the mid-section of the plastron (composed of the hyo- and
hypoplastra, covered by the abdominal scutes) is rigidly fixed to
the carapace by the bony bridge, but the front lobe of the plastron,
composed of the epiplastra covered by the single gular and paired
humeral and pectoral scutes, is moveable, as is the hind lobe com
posed of the xiphiplastra covered by the femoral and anal scutes.
The absence of the entoplastron in Kinosternon is almost unique
among turtles, and it is most interesting that the related genus
505
www.Ebook777.com
Mature male (right) and female Graptemys flavimaculata from the Pascagoula
River in Mississippi. Photographed at the Bronx Zoo.
Barbour’s map turtle, Graptemys barbouri, from the Chipola River in Florida.
Photographed at Ross Allen’s Reptile Institute.
506
Gulf Coast box turtle, Terrapene Carolina major. This is the largest and least
decorative North American box turtle. Photographed at Ross Allen’s Reptile
Institute.
Terrapene Carolina triunguis, the most westerly race of the common box tur
tle.
507
www.Ebook777.com
Staurotypus, which also has a hinged front plastral lobe, has a
well-defined entoplastron. It would appear that the hinges of these
two genera developed independently; a turtle with the normal ar
rangement of bones in a rigid plastron must either compress the
entoplastron from a diamond-shape to a triangle, or eliminate it
entirely, when it evolves a hinge between the epiplastra and the
hyoplastra. It is possible that the entoplastron of Kinosternon dis
appeared by a process of phylogenetic reduction and de-ossifica-
tion until it was absorbed into the hinge. In Staurotypus it appears
that the entoplastron has been moved forward and made triangu
lar rather than diamond-shaped, while the epiplastra have devel
oped postero-lateral projections, and the resultant lateral
vacuities have been infused with flexible cartilage.
All members of the family Kinosternidae have at least one
pair of barbels under the chin. Some mud turtles can seal their
shell openings, by raising the plastral lobes, as tightly as can a box
turtle, so that no part of the animal’s extremities remains exposed,
while others have relatively reduced plastra so that the shell can
not be tightly closed. Kinosternon leucostomum, scorpioides
cruentatum, abaxillare, and acutum have large plastral lobes
which allow complete closure, as do many specimens of integrum
and creaseri. In K. flavescens, K. hirtipes, K. sonoriense, and K.
scorpioides scorpioides, the plastron is large, but the flesh is still
visible when the lobes are raised. K. subrubrum and K. bauri have
smaller plastra, while K. herrerai, K. dunni, and K. angustipons
have very small plastra which provide very inadequate protection
for the soft parts.
The common mud turtle, Kinosternon subrubrum, is the most
familiar and most widespread representative of the genus in the
United States. It is a small species, usual specimens being only 3 or
4 inches in length, the record length being 4-7/8”. Three subspecies
exist: K.s. subrubrum, the eastern mud turtle, is found over a
large area of the eastern United States, extending from Long
Island, New York, south along the coastal plain to northern
Florida, east to western Mississippi, and north along the Missis
sippi Valley to southern Indiana, with an apparently isolated
colony in northwestern Indiana; K.s. steindachneri, the Florida
mud turtle, is found throughout the peninsula of Florida; and K.s.
hippocrepis, the Mississippi mud turtle, is found from southern
Alabama, extreme southern and western Mississippi, north to
southeastern Missouri, and south to southeastern Texas.
The eastern mud turtle has a rather low, brown shell with the
highest point well toward the rear, the posterior part of the cara
pace being strongly declivous. In old adults the carapace may
assume a slightly bilobed appearance. The plastron is yellowish,
brownish, or with markings of both colors, usually being some
508
what darker along the seams. The head is relatively large and the
jaws strong. The anterior lobe of the plastron is shorter than the
posterior lobe. Both lobes may be raised, but the sole parts of the
turtle always remain visible. The usual differences between the
sexes occur in this species, the male having a slightly concave
plastron and a longer, thicker tail with a rather distally located
vent; the tail of the male is normally kept curled to one side. The
soft parts are of an obscure dark shade, with light mottling on the
head; these light spots are sometimes vaguely aligned to form
broken stripes.
The eastern mud turtle is an aquatic species, but it may under
take overland migrations, usually during or after periods of heavy
rain. Archie Carr described the optimum habitat as perhaps repre
sented by a well-established, though fluctuating, shallow water
ditch with considerable aquatic vegetation. However, it has a high
tolerance for saline conditions and is often abundant on the inner
edges of tidal marshes and offshore islands.
The eastern mud turtle is reported to copulate in May and to
nest from the end of March to September, but most usually in
June. The female is said to start the nest with the forefeet, then to
turn round and finish the cavity—which may be up to five inches
deep—with the hind feet. Two to five eggs constitute a clutch, three
being the usual complement. The eggs are hard-shelled and elon
gate, variable in size but averaging about one inch in length and
0.58” in width. The hatchling is slightly less than an inch in length
and has a rather narrow shell; the carapace is almost black, but
the young are bright pink underneath, with a blackish figure con
centrated along the midline.
No systematic studies of the feeding habits of the common
mud turtle have been carried out; however, the diet is known to
include considerable quantities of insects and molluscs, and smal
ler quantities of crustaceans, amphibian larvae, and vegetation.
The animal could best be described as an opportunistic carnivore,
and in captivity it will eat almost any animal matter offered.
Newly captured specimens attempt to bite and may exude a stink
ing secretion, but both of these unpleasant habits are quickly out
grown in captivity.
The Florida mud turtle, Kinosternon subrubrum steindach-
neri, is found from Alachua County, Florida, south to Cape Sable;
intergrades between this race and subrubrum occur in northern
Florida and the Okefenokee Swamp. Dorsally, there is little to dis
tinguish the two races, but the plastron of steindachneri is
reduced, especially in the males, and the bridge is extremely nar
row—the width of the bridge is contained 2.5 to 3 times in the length
of the front plastral lobe, but only twice or less in subrubrum. Also,
steindachneri—males especially—is highly aggressive when
509
www.Ebook777.com
Florida box turtle, Terrapene Carolina bauri. The radiating stripes on the
high carapace are moderately distinctive. Photo by Dr. Herbert R. Ax
elrod.
The common box turtle, Terrapene Carolina Carolina, which ranges over much
of the eastern United States. Photo by Mervin R. Roberts.
510
Three-toed box turtle, Terrapene Carolina triunguis.
Terrapene coahuiia, a primitive aquatic box turtle from Cuatro Cienegas,

Coahuila, Mexico, Photographed at the Bronx Zoo.
511
www.Ebook777.com
handled and bites readily and hard; the most painful turtle bite I
ever sustained was dealt me by a freshly caught male of this race.
Sexual differences are the usual ones for the genus, with the
additional points that the heads of adult males often become stri
kingly enlarged, the bridge of the male is narrower, and the hind
plastral lobe shorter.
The Florida mud turtle is more aquatic than the northern race,
and its habit of remaining at the bottom of ditches and sloughs
gives the false impression that it is a much rarer animal. How
ever, individuals are picked up on the road crossing Paynes
Prairie, Alachua County, fairly frequently during very rainy
weather, and this turtle often invokes the ire of fishermen who
keep losing their bait to turtles instead of fish; such fishermen
near my house in Alachua County often lay mud turtles they have
caught out on the nearest highway to be crushed by passing cars.
Dietary studies of this turtle have not been carried out, but it is
possible that the diet includes a greater percentage of hard-shelled
organisms than does that of the eastern mud turtle, in view of the
heavier jaw surfaces, especially of the males. The eggs are laid
from March to mid-June in northern Florida; they average about
1.1” in length and 0.7” in width, and the normal clutch numbers
only two or three. The eggs are often not even completely covered
by the mother, but their hard shells prevent excessive dessication.
The Mississippi mud turtle, K.s. hippocrepis, is distinguished
from the eastern race by the presence of two well-defined light
Two distinct yellow stripes on the side of the head are characteristic of the
Mississippi mud turtle, Kinosternon subrubrum hippocrepis.
512
lines on each side of the head, as well as the usual light spots. In
size and shape there appears to be little to distinguish the forms,
although it is possible that the bridge of hippocrepis is relatively
wider than that of subrubrum. Carr recorded that this turtle
occurs in ponds, ditches, and shallow water generally. Conant
mentions it as a common turtle of bayous, lagoons, and great
swamps of the lower Mississippi Valley. Carr mentions an old
record of 45 individuals migrating in the same direction from a
shallow lake that was drying up near Waco, Texas.
The natural history of K. s. hippocrepis in Oklahoma has been
studied by I.Y. Mahmoud. Mahmoud found that the turtles are
active from early April until late October. During the summer
months, activity takes place primarily in the early morning hours
(4 to 9 a.m.) and the evening (4:40 to 10 p.m.). Maturity in females
is reached at 5 to 8 years of age, in males at 4 to 7 years. Carapace
length at maturity is 8 to 12 cm; growth takes place during some
what less than half the year—around 170 days. Stomachs of Okla
homa hippocrepis contained insects, crustaceans, molluscs,
amphibians, carrion, and aquatic vegetation. From tagging and
recapture experiments, Mahmoud concluded that the average
male has a home range of 0.12 acres, and the average female 0.13
acres. In suitable habitat the species may reach a density of 104.6
per acre. In three separate localities, Mahmoud found a rather
consistent sex ratio of approximately two males to three females
(47:71 at Cowan Creek; 16 to 24 at the Tishomingo Fish Hatchery;
16 to 28 in Lake Texoma).
The striped mud turtle, Kinosternon bauri, is a small species,
not exceeding 4.75” in carapace length, found throughout penin
sular Florida, and recent studies have shown that the species not
only also occurs in coastal Georgia, but also enters South Carolina,
having been found in bald cypress-tupelo gum swamps bordering
the Savannah River near Aiken. The shell is oval, usually relative
ly low, and slopes down more strongly posteriorly than anteriorly.
The unkeeled carapace varies from horn color to black, often
being red-brown, and there are almost always three longitudinal
light lines running the entire length of the carapace. The plastron
is moderately large but is too small to close the shell openings
completely even when the moveable lobes are fully raised. Little
pigment is visible in the plastron, although the edges of the scutes
may be somewhat darker than the central parts. The hind lobe is
the longest, while the central, fixed part is shorter than either lobe.
The axillary and inguinal scutes on each side usually have a nar
row line of contact. The interpectoral seam is very short or non
existent; it is not rare for the pectoral scutes to be quite widely
separated along the midline.
513
www.Ebook777.com
Florida diamondback terrapin, Malaclemys terrapin macrospilota.
Diamondback terrapin, Malaclemys terrapin, confined to brackish waters

along the eastern coast of the United States. Photo by Dr. Herbert R. Axelrod.
514
Florida chicken turtle, Deirochelys reticularia chrysea, a medium-sized turtle

with a very long neck.
Ornate turtle, Pseudemys scripta ornata, from Tortuguero, Costa Rica.
515
www.Ebook777.com
516
Kinosternon bauri. bauri
The skin of the head and limbs is dark grey to black; there are
two light stripes along each side of the head, the upper of which
reaches the snout. There are sickle-shaped scales on the forearms
and heels, and males, which are usually smaller than females,
have rough tubercular “clasping” areas on the femoral and tibial
sections of the hind limbs.
Two subspecies have been described. K. bauri bauri, from the
lower Florida Keys (Key West, Stock Island, Big Pine Key, and
possibly associated small islands), has an intensely pigmented
carapace and head, with the white stripes often almost obliterated
and an unstreaked or only slightly streaked mandibular beak. K.
bauri palmarum, from the Upper Keys north to Leon County,
Florida and the Savannah River, sometimes (particularly in the
southern parts of the range) has such reduced carapace pigment
that the bony sutures are visible in the live animal, but usually has
a rather heavily streaked mandibular beak.
This is a rather less aquatic species than most members of the
genus, and although it commonly lives in quite deep water, it is
also found in shallow or temporary bodies of water or even on dry
land. The eggs are laid from April to June in holes in sand or in
piles of dead water hyacinths. The eggs normally number one,
two, or three and are between 1” and 1.2” in length and 0.6” and
0.7” in width. A hatchling found crossing a road in the Everglades
on September 15 was 0.95” long and 0.82” wide.
517
www.Ebook777.com
Hatchling hieroglyphic turtle, Pseudemys concinna hieroglyphics, from the
south-central United States. Photo by Dr. Herbert R. Axelrod.
Yellow-bellied turtle, Pseudemys scripta scripts, a medium-sized deep-

shelled turtle from the southeastern United States. Photographed at Ross
Allen’s Reptile Institute.
518
Hatchling Pseudemys scripta callirostris from Colombia. Photo by Dr.

Herbert R. Axelrod.
Plastral view of Pseudemys scripta callirostris. Photo by Dr. Herbert R.

Axelrod.
519
www.Ebook777.com
Dietarily, the striped mud turtle appears to be an omnivorous
scavenger. Captive specimens will eat most foods offered them,
while stomachs of wild specimens examined by Einem were found
to contain seeds of cabbage palm, algae, small snails, insect frag
ments, beetle elytra, small bone fragments, vertebrae, juniper
leaves, and other vegetable debris. The disposition of freshly-
caught specimens varies greatly, some biting on the slightest
provocation while others seem to be very gentle. They can live a
long time in captivity.
The striped mud turtle is not usually subjected to such cold
winters that it has to undergo actual hibernation, but during cold
spells specimens have been found buried in piles of moist decaying
vegetation.
Kinosternon flavescens, the yellow mud turtle, is found in
most of Texas, Oklahoma, and Kansas, with the exception of the
extreme eastern parts of these states; it also occurs in extreme
southern Nebraska, eastern Colorado, eastern and southern New
Mexico, extreme southeastern Arizona, and perhaps southern
Utah. The range extends into northern Mexico, having been found
in the northern parts of all the states from Sonora to Tamaulipas,
but exactly how far it reaches into Mexico has not been deter
mined. Distinct colonies, referable to a separate subspecies
(spooneri), occur in certain stretches of the Illinois and Mississip
pi Rivers in Illinois, extreme northeastern Missouri, and extreme
southwestern Iowa. Furthermore, certain specimens from widely
separated points in Mexico (from Quitobaquito, 12 miles west of
Sonoyta, Sonora; Pedricena, Durango; and Jaral, Coahuila) have
been referred to a third subspecies (stejnegeri).
The yellow mud turtle has an oval, relatively flat shell of a uni
form olive-yellow to dark brown color with black seam borders. A
characteristic feature of this species is the elevation of the anteri
or part of the ninth marginal scute on each side in all but the smal
lest specimens. The posterior part of the tenth marginal is also
elevated, as in most mud turtles. The plastron is fairly wide, but
not large enough to close the shell openings completely. In color
the plastron is predominantly yellow, but the areas of recent
growth may be brown and the bridges and submarginal areas are
usually brown. The anterior and posterior parts of the plastron are
approximately equal in length and slightly longer than the fixed
middle section. The two hinges are well developed, and in adult
males there may also be a median line of flexure between the ab
dominal scutes. Males differ from the females in having slightly
concave plastra, long thick tails with a conical spur on the tip, and
in having opposed tubercular patches on the femoral and tibial
parts of the hind limbs. The head is relatively small for a mud tur
tle, and the upper jaw may be hooked or unhooked. The skin is mid-
520
grey in color with little indication of variegation; a variable patch

of yellow is present under the chin.
The yellow mud turtle reaches an adult length of four to five
inches; the record length for the species is 6 3/8 inches. It appears
that Illinois specimens may reach a slightly larger adult size than
those of the western race. Otherwise, the races are distinguished
by rather feeble characters; spooneri has, on average, a longer
gular scute (about half the length of the anterior lobe of the plas
tron), and overall darker coloration, with less yellow on the throat,
while stejnegeri has a gular almost 2/3 the length of the anterior
lobe of the plastron, a wide nuchal (about 1/4 the length of the
bridge), and a relatively long interfemoral seam. The Illinois race
is also reported to have a strikingly dark juvenile coloration, with
irregular but discrete light markings under the chin, which dis
appear at a shell length of about four inches.
This is a fairly aquatic species inhabiting muddy pools, but it
is also found in artificial habitats such as cattle tanks, ditches, and
even sewer drains. In wet weather or when the ponds dry up, it
may be found on land. Feeding habits in the wild have not been
A relict of an earlier prairie fauna in southern Illinois, Kinosternon flavescens

spooneri is barely distinguishable from typical K. flavescens. Photo courtesy
Illinois Natural History Survey.
www.Ebook777.com
Pseudemys malonei, found in freshwater ponds on Great Inagua Island,
Bahamas.
Half-grown red-eared turtle, Pseudemys scripts elegans. This was the most
common pet turtle in the United States. Photo by Dr. Herbert R. Axelrod.
522
Plastral view of red-eared turtle, Pseudemys scripta elegans. Photo by pr.

Herbert R. Axelrod.
Peninsular turtle, Pseudemys floridana peninsularis, a large, high-shelled

cooter from peninsular Florida. Photographed at Ross Allen’s Reptile In
stitute.
523
www.Ebook777.com
studied, but in captivity it will eat almost any animal matter of
fered. The normal egg complement of the typical race may be as
few as two, but spooneri lays 3 to 5 eggs. The eggs are a little over
an inch in length and about 0.6” in width.
This species is not particularly truculent when handled, but
freshly caught specimens exude a strong and rather nauseating
smell.
The isolated colonies of the yellow mud turtle in Illinois and
adjacent areas constitute, to quote Philip Smith: “One of the stri
king examples of a relict xerothermal period animal. The closest
relative occupies the Great Plains from eastern Kansas westward.
As a result of the climatic changes occurring in the prairie
peninsula, the race spooneri has evidently been reduced to a few
small remnant colonies.”
A recent analysis by Fred Punzo of the stomach contents of 37
yellow mud turtles from 4 km. east of Sanford Dam, Hutchinson
County, Texas, revealed that the species feeds in both aquatic and
terrestrial habitats. Diurnal feeding was usually aquatic, and at
this time flatworms, roundworms, annelids, crustaceans, aquatic
insect larvae and adults, snails and larval amphibians were eaten.
However, in the early morning and evening, when temperatures
are lower, the turtles forage on land and consume earthworms,
millipedes, centipedes, spiders and insects.
The Sonoran mud turtle, Kinosternon sonoriense, has the kind
of fragmented range that one expects of an aquatic animal that
lives in a predominantly arid area; it is found in ponds, springs,
creeks, and waterholes of intermittent streams in certain areas of
the southwestern United States and northwestern Mexico. In Ari
zona the species is known from the Gila River drainage in the cen
tral and southeastern parts of the state, and there is an isolated
colony at Quitobaquito Spring, Pima County, on the Sonora border.
The species is also found in southwestern New Mexico and ex
treme western Texas, and the range extends over undetermined
areas of northern Sonora, Chihuahua, and possibly Coahuila. In
California there are some old records for the Imperial Valley
(Palo Verde and Yuma Indian Reservations), but it is not certain
that these colonies still exist.
Over much of its range the Sonoran mud turtle is broadly sym-
patric with the yellow mud turtle, K. flavescens, but there is eco-
logic separation at least in some areas, sonoriense being more typ
ical of elevated country than flavescens. K. sonoriense is slightly
larger than flavescens, adults being up to about 6.5” in length, and
is easily distinguished from the latter species by the more elongate
olive or brown, often speckled and sometimes tricarinate, cara
pace which shows no elevation of the anterior part of the ninth
marginal scutes. The head lacks the slight supraorbital ridges
524
seen in flavescens, and the head and neck are heavily mottled with
white. The plastron is fairly large but too narrow to close the shell
openings completely. There is a shallow notch between the anal
scutes. The three sections of the plastron are of approximately
equal length. The gular scute is large and wide but slightly less
than half the length of the anterior lobe of the plastron. The sexes
are distinguished by the usual characteristics of the longer,
heavier, spur-tipped tail and the slight plastral concavity of the
male. The male has the usual areas of small rough scales on the
hind limbs. The female may reach a larger size, on average, than
the male.
The Sonoran mud turtle is found in a great variety of aquatic
situations, ranging from stock tanks and reservoirs to streams and
major rivers, though it does not occur in temporary bodies of
water nor in swift-flowing rivers. In Arizona it may be found at ele
vations of up to 5,500 feet, and up to 7,000 feet in the Gila Mountains
in New Mexico. The preferred habitats are small desert streams
and shallow ponds with attached submergent vegetation.
This species tends to adopt a nocturnal regimen during the hot
test part of the year, especially at lower elevations, but is diurnal
in spring and fall. In the Lower Sonoran desert the species appears
to be active year-round, but at higher elevations activity usually
begins in early March and ends in October. This is a highly aquatic
species which is normally found on land only when nesting. Locally
high population densities may be reached; A.C. Hulse estimated
that 325 animals were present in less than an acre of water in Tuley
Stream, Yavapai County, Arizona. The species is preferentially
carnivorous, a wide variety of animal remains having been found
in stomachs including Anisoptera, Physa, Trichoptera, Diptera,
Coleoptera, Ephemeroptera, Hemiptera, Zygoptera, Megaloptera,
ostracods, fish, and frogs; however, in situations where a high den
sity of turtles co-exists with a relatively low invertebrate fauna,
the diet may include greater amounts of plant matter, including
angiosperms. Chlorophyta, and Chara.
The reproduction habits of the Sonoran mud turtle have not
been described.
Kinosternon leucostomum, the white-lipped mud turtle, has an
extensive entirely tropical range, having been recorded from cen
tral Veracruz, Mexico, through northern Guatemala and Atlantic
drainages of Central America to northwestern Colombia. In
Mexico it extends inland as far as eastern Puebla and northern
Oaxaca and Chiapas. It does not reach the northern part of the
Yucatan Peninsula, but has been recorded as far north as Felipe
Carillo Puerto, Quintana Roo, and Becan, Campeche. The species
crosses into the Pacific drainage in the Golfo Dulce region of Costa
Rica and extends into Pacific Panama and Colombia to Ecuador.
525
www.Ebook777.com
Florida red-bellied turtle, Pseudemys nelsoni.
Suwannee River turtle, Pseudemys concinna suwanniensis, from Manatee

Springs, Florida.
527
www.Ebook777.com
In Mexico it is usually known by the name ‘pochitoque,’ in Colom
bia as ‘tabaculo,’ ‘culitap,’ or ‘morrocoy.’ It is a medium size spe
cies; Leger found that in Panama males become sexually mature
at about four inches, females at about 3.2 inches. Legler’s largest
Panamanian specimen was 6.8 inches long. Medem’s series of ten
males and 18 females from western Colombia showed respective
maximum lengths of 5.75” and 5.3”. The carapace is dark brown
moderately or even steeply elevated but not strongly declivous
posteriorly, and is unkeeled or has only a median keel. Old adults
often have a shallow median groove along the top of the carapace.
The plastron is large, and in half-grown and adult examples the
lobes are sufficiently developed to close the shell openings com
pletely. The front and hind plastral lobes are both distinctly longer
than the mid-section of thef plastron. The gular is fairly large but
normally less than half the length of the anterior plastral lobe. The
axillary and inguinal scutes are sometimes in contact, other times
separated; in fact this condition is so variable that those with pos-
eriorly-located inguinals have been described as a separate spe
cies (postinguvnale). The plastron and undermarginals are light
yellowish in color. There is no notch between the anal scutes. The
head is of medium size, and the upper jaw is virtually unhooked.
The jaws and underside of the neck are white, but the top and sides
o the head are dark brown except for a strong white or buff band
extending backward from the upper corner of each orbit. Fre
quently these light bands are infused with small brown spots.
There are three curved, sickle-like scales on the front face of each
forehmb and others on the heels. The soft parts are brown above,
uff below. Males have the usual longer, thicker tail but lack the
patches of homy tubercles found on the hind limbs of males of
most Kinosternon species.
The white-lipped mud turtle is an abundant species throughout
most ot its range, and despite its small size it is an important food
item in tropical Mexico and in Nicaragua. Its habits have not been
thoroughly studied, and we are indebted to Legler for most of the
information given below.
In Panama this species is amphibious rather than strictly
aquatic, individuals of both sexes frequently being found wander
ing about on land. It is omnivorous in diet, stomachs containing
comparable amounts of plant matter (Elodea and grasses) and
animal material (snails and insects). Reproduction occurs
throughout almost all of the year except for a short quiescent
period around January. The eggs are hard-shelled and elongate, as
is usual for mud turtles, and measure between 1.4” and 1.6” in
length and about 0.8” in width. Usually only a single egg is deposi
ted at a time, but five out of 21 clutches examined contained two
eggs. No proper nest is made, the egg(s) either being semi-buried
528
in a shallow cavity or simply covered with leaf litter. Three eggs,

hatched under artificial conditions but at normal environmental
temperatures, took between 126 and 148 days to hatch. In Colom
bia Medem found that K. leucostomum (recorded under the name
K. 'spurrelli) is nocturnal, hiding under stones and logs under
water during the daytime; it is found in brackish water in the
mouth of the San Juan River.
The species Kinosternon spurrelli was originally described by
Boulenger in 1913 on the basis of a single adult male from Pena
Lisa Condoto, Colombia. Subsequently a series was collected at
Pizarro, Choco, Colombia. The species was considered valid by
Medem in his monograph on reptiles of the Choco and by Mertens
and Wermuth in their 1955 and 1961 checklists of turtles of e
world though John Legler assumed it to be synonymous with K.
leucostomum. The only light I can shed on the question is based on
a series of six specimens of uncertain but presumably local origin
that I purchased in the market in Guayaquil, Ecuador. These tur
tles differed from typical leucostomum in being of consistently
smaller size despite the development of secondary sexual fea
tures, the largest being less than 4.5 inches long, and in having a
consistent pattern of numerous golden-yellow spots on the top of
the snout and along the buff-colored postorbital bar on each side.
These characters were not mentioned in the description of the type
specimen of spurrelli; the species was based on the dark plastral
pigmentation, a feature which is certainly not constant for the
Colombian population. Nevertheless, I feel that the Ecuadorian
and Central American populations are subspecifically separable,
and that the type of spurrelli probably represents some sort 0
intermediate between these populations. I believe therefore that it
would be appropriate to designate the coastal Ecuadorian mud
turtles Kinosternon leucostomum spurrelli.
The first mud turtle to be introduced to science was the scor
pion mud turtle, Kinosternon scorpioides\ the common and scien
tific names refer to the homy spur-like tail-tip of the male, though
the function of this spur is to hold the female during copulation
rather than to inject venom. The scorpion mud turtle is found over
much of tropical South America, having been recorded from many
parts of the Amazon drainage in Brazil, Peru, and Ecuador, as
well as in the north-draining rivers in Colombia, Venezuela, and
the three Guianas. To the south it reaches as far as Bolivia and
northern Argentina. Four subspecies of this species in South
America have been described: scorpioides, occupying most of the
range of the species; pachyurum in Bolivia; seriei in northern Ar
gentina' and carajasensis from the isolated elevated plateau of
Serra dos Carajas in southern Para, Brazil. These subspecies are
not strongly characterized, and further studies may well prove
529
www.Ebook777.com
Hatchlings of Pseudemys scripts dorbignyi, a form from southern Brazil and
northern Argentina. The range of this subspecies is widely separated from
the ranges of other members of the species. Photo by Harald Schultz.
Fruit-eating terrapin, Pseudemys felis , restricted to Cat Island in the

Bahamas. Photographed at the London Zoo.
530
European pond turtle, Emys orbicularis, a variable species from southern

Europe, North Africa, and the Middle East. This individual is from the Caspian
Sea region of Iran.
Blanding’s turtle, Emydoidea blandingi, from the Great Lakes region of

Canada and the United States. Photo by Dr. Herbert R. Axelrod.
531
www.Ebook777.com
Kinosternon scorpioides scorpioides, the scorpion mud turtle. This
specimen is from Trinidad. Photo by R.J. Church.
them invalid. However, pachyurum is supposedly distinguished by

having lower and less distinct keels than scorpioides; carajasensis
by having a rather elevated, tectiform carapace with almost no
trace of the lateral keels, a vestigial nuchal scute, and a relatively
large head; seriei by the following combination of characters:
width of nuchal equal to its length; interanal seam 3 to 4 times
length of interfemoral seam; cartilaginous infusion present be
tween abdominals and femorals; extremities grey in color; and
overall size greater, and build more massive, than in scorpioides.
The scorpion mud turtle reaches an adult length of five or six
inches. It has an elongate dark brown carapace with narrow cen
tral scutes and three strong keels in all except old adults. The
widest point of the shell is often well posterior to the mid-point. The
plastron is moderately large but is too small to close the shell open
ings completely, and there is a shallow notch between the anal
scutes. The front and middle sections of the plastron are of approx
imately equal length, but the hind lobe is distinctly longer. The
axillary and inguinal are usually in contact. Males have a barely
discernible concavity centered on the middle of the posterior hinge
and also have more intensely marked heads, the usual pattern in
males being one of bold white streaks on an almost black back
532
ground. The top of the head is uniformly dark brown or black in

males, while the throat is mottled with light markings. The head of
the female, on the other hand, does not have such strongly contras
ting markings. Males have stronger throat barbels than females,
and also have the usual longer, thicker tails, with the vent more
distally located. The strong sexual dimorphism in tail length led
the Brazilian explorer Spix to apply different scientific names to
the males and females (Kinosternon longicaudatum, and K. brevi-
caudatum, respectively). On the other hand, males lack the horny
thigh tubercles found on hind legs of most Kinosternon species.
Despite the wide range of the species, the habits of the scor
pion mud turtle are poorly known. The eggs, probably usually be
tween one and three in number per nest, are about 1.6” long and
about 0.75” wide. The courtship, observed by Owen Sexton for
Venezuelan specimens, involves the male nibbling at the edge of
the female’s carapace. The copulatory position involves the male
holding on to the shell of the female with all four feet as well as the
hooked tail. During copulation the neck of the male is fully exten
ded and bent downward toward that of the female. After intromis
sion, the female dislodges the male by moving about rapidly and
biting his head.
The red-cheeked mud turtle, Kinosternon scorpioides cruen-
tatum, is usually considered a separate species, but in fact it inter
grades with the scorpion mud turtle in Panama. The name K.s.
panamense has been used for such intermediates, but it is prob
ably more accurate simply to consider Panamanian specimens as
intergrades between cruentatum and scorpioides. K.s. cruentatum
is found in Pacific drainages of Costa Rica, Nicaragua, Honduras,
El Salvador, Guatemala, and the States of Chiapas and Oaxaca in
Mexico. It extends across the Isthmus of Tehuantepec to the Gulf
drainage in Tabasco, east as far as Stann Creek District, Belize. It
reaches right to the northern end of the Yucatan Peninsula, there
being records from Dzibalchen, Isla del Carmen, Laguna Silvituc,
and Xpujil in Campeche; Tizimin, Progreso, and Puerto Telchac
in Yucatan; one mile west of Puerto Morelos in Quintana Roo; and
Cozumel Island, off the coast of Quintana Roo. There is also an iso
lated population in southern Tamaulipas and eastern San Luis
Potosi, separated from the nearest colony to the south (in
Tabasco) by about four hundred miles. Locality records exist for
Sierra del Bernal, 5 miles east of Magiscatzin, Tamaulipas; and
near Ebano and at Tamuin, San Luis Potosi.
The red-cheeked mud turtle typically differs from the scorpion
mud turtle in having a broader, more humped shell with a com
pletely closeable plastron with no interanal notch, and usually with
red or orange spots on the side of the head. The carapace may be
yellow, orange, or black; specimens with light shells usually have
533
www.Ebook777.com
Muhlenberg’s turtle, Clemmys muhlenbergi, from Pennsylvania and states

adjoining to the east. This is the smallest and perhaps rarest North American
turtle.
Pacific pond turtle, Clemmys marmorata, from California. Photo by Dr. C. M.

Bogert.
535
www.Ebook777.com
relatively dark scute edges. Typically the carapace is tricarinate,
but the keels are lost in very old individuals, and all the specimens
I have seen from Pacific Guatemala and Honduras had no trace of
dorsal keels. The plastron and under-marginals are usually orange
with dark scute edges. The gular may be more or less than half the
length of the anterior plastral lobe. The interpectoral and inter-
femoral seams are shorter than is typical of the scorpion mud tur
tle. Both moveable lobes are a little longer than the fixed mid-sec
tion of the plastron. The axillaries and inguinals may be in contact
or may be widely separated. The soft parts are grey in color; the
male has the usual long, thick tail but normally lacks rough ‘clasp
ing organs’ on the hind limbs (though these were reportedly dis
tinct in a specimen from San Luis Potosi). The shell is often very
thick, especially in the more northern parts of the range. Adults
are five or six inches in length.
In Chiapas, the red-cheeked mud turtle (locally known as ‘cas-
quito amarillo’) is reported to live in rivers and streams, occasion
ally being seen basking on emergent rocks. The female makes her
nest close to the water and lays about ten eggs ih March or April,
according to Alvarez del Toro.
Mud turtles are very abundant on the island of San Andres in
the western Caribbean, though serious drought apparently deci
mated this isolated population in 1972. These turtles have been
named a separate subspecies (albogulare). In fact, they are very
close to typical cruentatum, though they may, on average, have
less pigmentation on the jaws and throat. A further subspecies,
consors, reported from northern Yucatan and Cozumel Island,
does not appear to be valid.
At the northern limit of its range, K.s. cruentatum may or may
not intergrade with K.s. integrum (or K. integrum). In view of this
uncertainty, it may be best for the present to use the binomial.
Kinosternon integrum has been recorded from the Pacific coastal
State of Mexico from Sonora to Oaxaca, as well as the largest of
the Tres Marias Islands (Maria Madre). Inland, it has been recor
ded from Morelos, Aguascalientes, Guanajuato, San Luis Potosi,
and Puebla. It also reaches Veracruz, though how near it ap
proaches the Gulf Coast has not been determined. This is a some
what nondescript and highly variable species. The carapace is un
keeled in adults, though juveniles may show median and even lat
eral keels, and is more strongly declivous behind than in front. The
central scutes (except for the first) are wider than long. The over
all coloration of the carapace is brown or black, though some spe
cimens are relatively light brown with numerous small black
spots. The plastron is yellow, often with black scute edges, and is
almost but not quite large enough to close the shell openings com
pletely. The interfemoral seam is relatively long, and the gular is
536
broad and approaches half the length of the anterior plastral lobe.
The fixed mid-section of the plastron is shorter than both the an
terior and posterior lobes. The dorsal surfaces of the head and soft
parts are mid-grey with small buff spots on top of the head. The
jaws and soft undersurfaces are white with a few dark grey spots
and streaks on the jaw surfaces and sides of the head. Males do not
have clasping organs on the hind limbs.
To the north, the range of integrum reaches suspiciously close
to that of sonoriense (in southern Sonora), and the species are
somewhat similar morphologically. However, intergradation has
not been demonstrated. It might be thought that the presence of
clasping organs in male sonoriense and their absence in integrum
would be a strong character for separating the species, however,
clasping organs in some species (e.g. leucostomum and scorpioi-
des cruentatum) may be totally absent or partially or even fully
developed. The precise use of these organs in relation to hind limb
morphology, plastral width, and general ethology has not been
studied; it would be most interesting to know more about them.
Certain mud turtles from the state of Veracruz appear to show
characteristics of both integrum and leucostomum', they have
massively heavy, domed, unkeeled, dark brown carapaces and
yellow plastra which are large enough to close the shell openings
completely. The plastron has a short mid-section with much longer
anterior and posterior lobes, and the head is nearly uniformly
dark, with heavy dark streaking on the jaws. Whether these are
genuine intergrades, local variants of one or other of the two spe
cies mentioned above, or representatives of an undescribed spe
cies has yet to be determined.
Kinosternon integrum is for the most part a lowland coastal
form. It is replaced in the main Mexican plateau, from Chihuahua
southward to the state of Mexico, by the rather similar Kinoster
non hirtipes hirtipes; records for this form exist for the states of
Chihuahua, Sinaloa, Michoacan, Colima, Guanajuato, San Luis
Potosi, Hidalgo, Mexico, and the Distrito Federal.
Kinosternon h. hirtipes has a dark brown, rather elongate
shell, without keels, with the widest point near the middle, and the
highest point at the rear of the third central scute. The central
scutes are broader than long, and the tenth marginals are greatly
elevated posteriorly. The plastron, which is yellow but often
strongly infused with brown or black, is just too small to close the
shell openings completely. The two moveable lobes are each
slightly longer than the fixed mid-section of the plastron, and the
posterior hinge is slightly convex. There is a very small interanal
notch. The gular scute is normally a little less than half the length
of the anterior lobe. The head is rather broad, with the upper jaw
of the male slightly hooked. The head and neck are dark brown
537
www.Ebook777.com
Heosemys grandis from Indonesia.
Mauremys mutica. Photo by Dr. C.M. Bogert.
538
Caspian terrapin, Mauremys caspica caspica, from central Iran.
Spanish terrapin, Mauremys leprosa. This species is plentiful in Spain and

Algeria. Photo by G. Marcuse.
539
www.Ebook777.com
with numerous light spots, which may become confluent. The male
has a homy spur on the tip of his tail and also roughened clasping
scales on the inner surfaces of the hind limbs—a point of differen
tiation from integrum which gives this species its name.
Kinosternon h. hirtipes is sold in numbers in certain markets
in Mexico City, apparently for aquaria rather than for food. In Chi
huahua, Legler reports that four to seven eggs are laid at a time.
The hatchlings of hirtipes are virtually identical to those of subru-
brum, even to the extent of having an orange or pink plastron with
a dark central area.
The other subspecies, Kinosternon hirtipes murrayi, is found
in parts of northern Durango, along the Rio Conchos in Chihuahua,
and just enters the United States at Presidio County, Texas. In Chi
huahua it is reported to be equally abundant in mountainous and
lowland areas. This subspecies is defined by the larger gular (half
or more of the length of the anterior plastral lobe), relatively
longer interpectoral and interfemoral seams, and shorter inter-
humeral seam. It is also possible that the head is more lightly
colored than in the typical race.
Legler reports that K.h. murrayi may lay four or five eggs at a
time, about 1.1” in length by 0.6” wide.
While it is possible that murrayi may intergrade with sonori-
ense in west Texas, or hirtipes may intergrade with integrum in
Mexico, neither possibility has yet been demonstrated to be the
case.
Kinosternon abaxillare is a highly localized species apparent
ly restricted to the central depression of the state of Chiapas,
Mexico, in which the state capital, Tuxtla Gutierrez, is located. It
is locally known as ‘casquito pardo’ which may be literally trans
lated as ‘little brown box,’ to distinguish it from the ‘little yellow
box’ (Kinosternon scorpioides cruentatum). It is a medium-sized
species, up to about six inches in carapace length, with a rather de
pressed shell, strongly declivous behind and with traces of three
keels in all except old adults. The carapace is brown, the plastron
yellowish with brown infuscations. The species derives its name
from the complete absence of the axillary scutes, a condition found
in other mud turtles only as an occasional variant. The inguinal
scutes, on the other hard, are well-defined, elongate, and relative
ly wide in the area of the posterior hinge. The plastron is large, and
the raised lobes completely close the shell openings except for a
narrow opening between the nuchal area and the gular scute of the
raised front lobe. The posterior margin of the abdominal scutes is
remarkably convex; it would appear that a curved line of flexure
would impair the function of the hinge, but in fact the hinge is just
as effective as the perfectly straight posterior hinge in the plastron
of the sympatric K.s. cruentatum. The fixed mid-section is the
540
Restricted to Chiapas, Mexico, Kinosternon abaxillare is very closely related

to Kinosternon scorpioides.
541
www.Ebook777.com
Rhinoclemys funerea from Tortuguero, Costa Rica. A predominantly black,
highly aquatic species that sometimes reaches a length of 18 inches.
542
Rhinoclemys pulcherrima manni, a small semi-aquatic turtle from Nicaragua.
Rhinoclemys pulcherrima , a beautifully marked turtle from southern Mexico.
543
www.Ebook777.com
longest part of the plastron, while the front moveable lobe is the
shortest. The gular scute is less than half the length of the front
lobe. There is no notch between the anal scutes. The hind lobe of
the plastron widens considerably posterior to the hinge.
The head of K. abaxillare is fairly small, but the beak is
pointed. The head is olive-brown in color with extensive light yel
low spots and rays. The soft skin and the limbs are brown.
This species has been observed in the wild by Miguel Alvarez
del Toro, Director of the Tuxtla Gutierrez Zoo. Del Toro records
that the casquito pardo is found in streams and marshes, not des
pising even the most insignificant bodies of water. In the dry sea
son it hides itself under rocks or buries itself in holes in the banks
of streams or in the mud at the bottom. In captivity it prefers meat
and is easily caught on baited hooks. Six to twelve eggs are laid at
a time, the season extending from March to May, and the hatch
ling turtles emerge about three months later. This circumscribed
breeding season and large clutch size are in striking contrast to
what we know of the habits of most other tropical Kinosternon spe
cies.
Kinosternon creaseri is a rather poorly-known species of mud
turtle from the northern, relatively arid part of the Yucatan Pen
insula of southern Mexico; the type series, consisting of an adult
male, three adult females, and four juveniles, was collected by Ed
win Creaser one mile south of the Hacienda at Chichen Itza, Yuca
tan. Since then, William Duellman has obtained fifteen further
specimens—seven from Piste, near Chichen Itza; six from Pueblo
Nuevo X-Can, Quintana Roo; one from Champoton, Campeche;
and one from Dzibalchen, Campeche. To these localities Smith
and Taylor add Vivienda de Platanal, Quintana Roo. The turtles
are confined to the cenotes, or water-filled limestone sinkholes,
which are dotted over the northern part of the peninsula. Whether
the various populations are completely isolated from each other,
or whether they migrate from one cenote to another, either over
land or through underground streams, is unknown. Kinosternon
creaseri is a medium-size mud turtle reaching a maximum cara
pace length of about five inches. The carapace is normally dark
brown, with the highest point well to the rear, and the posterior
part strongly declivous. Hatchlings and juveniles have three keels,
but in adults only the median keel remains. The plastron, which is
long and wide but just too small to close the shell openings com
pletely, is yellowish-brown with darker seams and is smoothly
rounded behind, with no median notch. The anterior lobe of the
plastron is always longer than the fixed middle section, and in
adults the gular scute is more than half the length of the anterior
lobe. In some juveniles, however, the gular may be a little less
than half the length of the front lobe. The head is large, especially
544
in the male, and the upper beak is strongly hooked, the hook being
flanked by notches that become deeper with age. The posterior
border of the single scale that covers the front part of the head ex
tends from orbit to orbit. The head and neck are normally black
above with barely-discernible lighter dots. The sides of the head
are somewhat lighter, and the underside is whitish-grey with black
streaks and specks on the throat. The jaws are brownish with fine
darker streaks.
Two specimens of Creaser’s mud turtle found by Duellman
had pale tan carapaces, pale creamy-yellow plastra, and pale
greyish-brown limbs and heads. They were found in a solution
cave in the limestone, and their pallid coloration may indicate a
prolonged sojum in virtually lightless waters.
In the more humid areas to the south, K. creaseri is replaced
by K. acutum, whose range extends from central Veracruz
through Tabasco and Campeche to northern Guatemala and
Belize. This is a small species, not more than 4.5” in carapace
length, distinguished from K. creaseri by the smaller head, com
pletely closeable shell, and relatively shorter anterior plastral
lobe—the anterior lobe is the same length as, or shorter than, the
fixed mid-section of the plastron. This species agrees with K.
creaseri in the lack of a notch in the rear of the plastron and the
presence of a long gular scute more than half the length of the
anterior plastral lobe. The carapace is brown in color with a single
median keel and with the highest point well to the rear. The last
Plastral view of Kinosternon acutum, which ranges from southeastern Mex

ico to Guatemala. Photo by G. Marcuse.
www.Ebook777.com
Rhinoclemys punctularia from Paramaribo, Surinam.
Rhinoclemys punctularia from Guyana. Photo by Dr. Herbert R. Axelrod.
546
Rhinoclemys areolata, a small semi-terrestrial species from Belize.
Rhinoclemys rubida from Mitla, Oaxaca, Mexico. Photo by Walter S. Miller.
547
www.Ebook777.com
two marginal scutes and the supracaudals on each side are in
creasingly elevated, with convex upper margins. The plastron is
yellowish or brown with dark brown seams, constricted at the pos
terior hinge, and with the axillary and inguinal scutes either sepa
rated or only narrowly in contact. The pectoral scutes have an
appreciable common boundary, but not as long as that between the
humerals. The head is basically brown above and below, with light
mottling on the top and sides; this light mottling becomes more ex
tensive on the mandibular plates and on the underside. There is a
single pair of chin barbels. The eyes are often strikingly red, with a
black horizontal line through each, and other small irregular areas
of black pigment. The beak is slightly hooked in the female, more
strikingly so in the male. The limbs are pale grey-brown with
brown vermiculations on the upper side of the forelimbs. The tip of
the tail of the male is armed with a conical horny spur.
The habits of this species have not been studied. In some areas
(e.g. near La Libertad, El Peten, Guatemala), it occurs sympatri-
cally with both K. scorpioides cruentatum and with K. leuco-
stomum.
Although most of the tropical species of Kinosternon have
large plastra which provide good protection for the extremities,
three rather well-separated forms have strikingly small plastra
which leave considerable soft areas exposed even when the move-
able lobes are fully raised. These three species are Kinosternon
herrerai, K. angustipons, and K. dunni. All are of relatively recent
discovery, having been first described in 1925, 1965, and 1947
respectively, and all are still known from relatively few speci
mens.
Herrera’s mud turtle, Kinosternon herrerai, is found in east-
central Mexico, having been recorded along the coast from the
Municipio de Aldama, Tamaulipas, south to the region of Jalapa,
Veracruz, and inland into eastern San Luis Potosi, extreme nor
thern Puebla, and probably eastern Hidalgo. This is a relatively
large species, a maximum carapace length of about 6.8” being
reached; males reach a larger size than females, an unusual situa
tion in small turtles. The carapace is depressed, almost oval, but
slightly expanded posteriorly in females and young males. The
carapace is only moderately declivous behind. No traces of lateral
keels are present, but there may be indications of a median keel on
centrals 3, 4, and 5. Marginals 1 to 9 are very low, but the tenth is
elevated posteriorly and the supracaudals are elevated to this
same height (in most specimens) throughout their length. The first
central scute is narrow and is always well separated from the
second marginals. The plastron is much too small to close the shell
openings. The hind lobe has a deep median notch in adult males
but is only slightly indented or truncated in females. The axillary
548
Herrera’s mud turtle, Kinosternon herrerai, is a poorly known species from

eastern Mexico.
and inguinal scutes always have a considerable line of contact.

The gular is less than half the length of the front lobe of the plas
tron; its width is usually comparable to its length. Both lobes are
always longer than the middle, fixed section of the plastron. In
males the front lobe is the longer; in females the longer is usually
the hind lobe. The males have well-developed roughened clasping
surfaces on the inner faces of the hind limbs, and they also have a
slight concavity toward the rear of the plastron, and the mid-sec
tion of the plastron is set at an angle, rather than being more or
less parallel to the top of the carapace, as in females. The jaws are
strongly hooked, particularly the lower one.
In color the carapace is uniformly brown or olive. The ventral
part of the marginals and the plastron are immaculate pale yel
low, though sometimes stained with red-brown ferric oxide. The
top of the head is pale buff with numerous small black spots. The
jaws are pale, with feeble to strong vertical black stripes. The
outer surfaces of the limbs are dark grey, the inner surfaces white.
Two bony shells of this species examined by the writer each con
tained only four neural bones, but others may have five.
The habits of Herrera’s mud turtle have not been studied in the
field; several of the rather few specimens available with good
locality data were picked up crossing highways, and it is possible
that this species is less aquatic than some of its relatives. It is
rarely sold for food, but fair numbers are preserved, lacquered,
mounted (often in grotesque positions, holding miniature musical
549
www.Ebook777.com
Heosemys grandis from Southeast Asia. Photo by Mervin F. Roberts.
Ceylon terrapin, Melanochelys trijuga thermalis.
550
Three-banded box turtle, Cuora trifasciata. This species is more aquatic than
the American box turtles. Photographed at the London Zoo.
Malayan box turtle, Cuora amboinensis.
www.Ebook777.com
instruments), and sold to tourists, along with hawksbill shells and
blown toads dyed green, in street markets in Tampico and other
cities.
Kinosternon angustipons is found in a restricted area of the
Caribbean drainage of Central America from the mouth of the San
Juan River in Nicaragua through eastern Costa Rica to the vicin
ity of Almirante, Bocas del Toro, Panama. It appears to be a rare
species throughout its range. It is small, not exceeding 4.8” in car
apace length; females are, on average, slightly larger than males.
The carapace is relatively low, with the highest point toward the
rear, and is unkeeled. The tenth marginal scutes are elevated pos
teriorly, but the supracaudals are low. The plastron is very small;
the axillary and inguinal on each side are in broad contact. The
hind lobe is slightly longer than the front lobe, and the fixed mid
section of the plastron is shorter than either. The gular is short and
wide; the pectoral scutes have a strong median line of contact. The
sides of the hind lobe are rounded and there is a strong notch be
tween the anal scutes. The head is relatively small, the jaws rather
weak and not hooked, the snout long but rounded, and the limbs are
rather small. Males have much longer tails than females, with
blunt, soft tips lacking the horny spur found in the males of many
other species. The males also have more extensive areas of soft
skin in the plastral hinges and a deeper notch between the anal
scutes. Clasping organs (the usual patches of roughened scales)
are present on the hind limbs of the males. The ground color of the
limbs, tail, and dorsal part of the neck is pale grey to brownish
grey, slate in the darkest areas. The ventral surface of the neck is
yellowish cream with a small grey area just in front of the plastron
and with pink on the gular region. The horny jaw sheaths are yel
lowish cream grading into pale brown where they approach the
soft skin of the head.
This species is found in shallow swamps, usually being found
in conjunction with Rhinoclemys funerea, Chelydra serpentina
acutirostris, and Kinosternon leucostomum. From very sparse
data, it appears that the eggs, which are about 1.6” by 0.9” in size,
number from one to four and more than one clutch may be pro
duced annually.
In captivity, Kinosternon angustipons feeds readily on meat
and fish, but in the wild it appears to be an omnivorous, opportu
nistic feeder, though eating more vegetable than animal matter.
Kinosternon dunni appears to be closely related to K. angusti
pons, but differs in reaching a much larger adult size—in fact the
largest known shell, 7 inches long, makes this species possibly the
largest of the genus. Other differences include the relatively wider
bridge of dunni (about 24% of the length of the plastron, instead of
17 to 20%), the larger head with strong jaws, and the higher cara-
552
The very rarely col

lected Colombian
Kinosternon dunni.
Notice the bulbous
nose. Photos by Dr.
F. Medem.
553
www.Ebook777.com
Reeves’ turtle, Chinemys reevesi, a small species very plentiful in the Far
East. Photo by Dr. Herbert R. Axelrod.
554
Indian roofed turtle, Kachuga tecta. Photographed at the London Zoo.
Kachuga smithi. Photo by Dr. Sherman Minton.
555
www.Ebook777.com
pace; also, the shape of the plastron differs In the two species, the
sides of the hind lobe being smoothly rounded and noticeably con
stricted at the hinge in angustipons, less smoothly rounded and not
constricted at the hinge in dunni. Nevertheless the species are gen
erally very similar and share such features as the strikingly re
duced plastron and the bulbous rather than constricted snout,
especially in adult males. It is possible that they should be con
sidered subspecies only, but at the present time the ranges of the
two species appear to be separated by at least 400 miles, and there
is thus no evidence of intergradation.
Kinosternon dunni is either an extremely rare turtle or one
whose way of life makes it so rarely encountered that even the
local people are often unaware of its existence. Up to now, the spe
cies is only known from four preserved specimens (one male and
three females), and a single bony carapace, probably of a female.
The known specimens are all from the Departamento del Choco,
on the Pacific coast of Colombia; locality records are: Pizarro,
near the mouth of the River Baudo; Noanama, on the Rio San
Juan; Rio Pepe, a tributary of the Baudo; and Cano Sando, a tribu
tary of the Rio Pepe. Local people also report the existence of this
species in various streams draining into the Gulf of Tribuga, but
there is not even circumstantial evidence suggesting its occur
rence north of the Jurubida River, or south of Buenaventura.
Kinosternon dunni has a uniformly dark brown carapace with
a slight reddish tinge. The plastron In the available specimens is
nearly as dark as the carapace, but this may be due to staining.
The head is dark brown above in the male, with numerous small
yellow spots; the sides and underneath of the head are light yellow
ish. The neck is grey above, light grey with yellowish markings be
low. The limbs are light grey in color. The female is less strongly
marked than the male, with grey on top of the head and whitish
with light grey on the sides of the head and on the neck. The sexes
also differ in that the tail of the male is longer and thicker, and the
male also has ‘clasping organs’ on the hind limbs, composed of
rough but spatulate rather than pointed scales.
Kinosternon dunni appears to be a mollusc-eater, at least jud
ging by the preferences of the male specimen in captivity and the
contents of the cloaca of a female which died after several months
in captivity, reportedly without feeding. The eggs, which are prob
ably laid throughout the year in small batches of two or so, are the
largest known for the genus, being about 1.8” long and 1 inch wide.
Mud turtles in Mexico have been studied intensively in recent

years by Jim Berry and John Iverson, and at the time of writing
these authors have several new forms and revisions in press. A
new species, to be named Kinosternon alamose, is restricted to the
556
vicinity of Alamos, Sonora, and the lower Rio Yaqui, the range
extending essentially from half-way between Hermosillo and
Guaymas to Alamos. It is related to scorpioides, the males lacking
horny thigh patches; it has widely separated axillary and inguinal
scutes, and usually no contact between the first central scute and
the second marginals. It shows some superficial similarities to
flavescens, such as the broad unkeeled shell, but the ninth
marginal is not elevated and the plastron is relatively larger. The
latter is an adaptation to arid conditions, this species spending
much of the year in estivation, during which a large plastron helps
lessen evaporative water loss.
John Iverson has recently shown that the Mexican subspecies
of Kinosternon flavescens, K.f.stejnegeri, is indistinguishable
from the fossil K.arizonense', the race which occurs in extreme
southern Arizona and along the central part of Sonora is therefore
properly known as K.f.arizonense. An additional subspecies,
K.f. durangoense, has a limited distribution near the juncture of
the states of Chuhuahua, Durango, and Coahuila; it differs from
K.f.flavescens in the proportions of certain scutes, notably the first
central, the nuchal, and certain plastral scutes.
Another localized new species is to be called Kinosternon
oaxacae; it is apparently restricted to the vicinity of Pochutla,
Oaxaca, between the integrum population in the Rio Verde and the
cruentatum population in the Rio Tehuantepec. It is a relatively
large species (up to 6.3 inches in length) with a depressed, strongly
tricarinate carapace, a relatively small plastron, a distinct
posterior plastral notch (especially in males), relatively long
bridges and fixed mid-section of the plastron, axillary and inguinal
scutes in contact, inguinal in contact with marginal scute 5, first
vertebral scute contacting marginal 2, no patches of roughened
scales on hind limbs, and tails of both sexes terminating in horny
spines.
Other taxonomic changes contemplated by Iverson and Berry
include the designation of a new subspecies, K. hirtipes
chapalaense, in Lake Chapala, and relegation of Kinosternon
abaxillare to subspecific rank within K. scorpioides.
GENUS STERNOTHERUS
The musk turtles, genus Sternotherus, are closely related to
the mud turtles of the genus Kinosternon. They differ in that the
plastron is always small; the hind lobe, instead of having rounded
sides and a rounded or absent posterior notch, has convergent and
almost straight sides, and the posterior notch is angular. Also, the
interpectoral seam is long in musk turtles, usually longer than the
interhumeral seam, while it is usually (but not always) very short
or non-existent in mud turtles. In mud turtles the suture at the rear
557
www.Ebook777.com
Batagur baska, a large aquatic turtle from Southeast Asia. Drawing by Peter
Parks.
Hardella thurji from the Indus River in India. Photo by Dr. C. M. Bogert.
Siebenrockiella crassicollis from Southeast Asia. Photo by Mervin F.

Roberts.
an Asiatic species in which the female is much larger than the

Hardella thurji,
male. Photo by G. Marcuse.
559
www.Ebook777.com
of the hypoplastra is straight and coincides with the straight pos
terior seam of the abdominal scutes, forming a well-developed
hinge. In the musk turtles, the seam at the rear of the abdominals
forms an obtuse angle on the mid-line and does not coincide with
the suture at the rear of the hypoplastra, so that the hind lobe of the
plastron has very little flexibility. Also, the gular scute of musk
turtles is never large and is sometimes absent, while it is invari
ably present and sometimes half the length of the anterior plastral
lobe in the mud turtles. Finally, some musk turtle species have a
marked tendency to develop extensive areas of soft skin between
the plastral scutes; this does not normally occur in mud turtles.
Certain forms included within the genus Kinosternon, such as
K. subrubrum steindachneri and K. herrerai, would appear to be
intermediate between the two genera in some respects.
Three species are currently included within the genus Stemo-
therus; one of these (S. odoratus) is very widely distributed in the
eastern United States, while the other two are relatively localized
in the southern states.
The common musk turtle, stinkpot, stinking jim, or skillpot,
Stemotherus odoratus, is found from the coast of Maine, west
through southern Ontario to extreme southern Wisconsin, south to
Ventral view of a stinkpot, Stemotherus odoratus, showing the greatly reduc

ed plastron with large fleshy areas between the scutes.
560
the tip of Florida, and west to east-central Texas. There is also a

record from Presidio County, Texas, and one from Sauz, Chihua
hua. The latter, based on a single specimen collected in 1903, is the
only record of the genus Stemotherus from outside the United
States.
The common musk turtle is a small species, being mature at 3
to 4 inches and with a record length of 5 3/8 inches. Hatchlings are
tiny, about 3/4” to 1” long, with black, steeply humped shells with
three keels and a small white spot at the edge of each marginal
scute. As the turtle grows, the shell becomes proportionally lower
and more elongate; the keels are lost when maturity is reached.
Also, the black color may become paler, sometimes revealing a
spotted pattern, but the marginal spots disappear. The plastron,
which includes a gular scute, has a symmetrical black pattern in
the hatchling, but this is soon lost. The bridge is quite narrow, and
the axillary and inguinal scutes on each side are in contact.
The head of the common musk turtle is unspotted and black or
grey in color with two white lines on each side extending backward
from the tip of the strongly projecting snout; one of these lines
passes above the eye, the other below. There is a pair of barbels on
the chin and another pair on the throat. The neck is long and highly
flexible, and since the animal can bite very hard it is wise to
handle it only by the rear third of the shell. The limbs are relative
ly small and have the curved, sickle-shaped scales on the outer
surface of the front limbs and the heels that are present in many
members of this family. Males differ from females in having
longer, thicker, spur-tipped tails, shorter plastra with more exten
sive areas of soft skin between the plastral scutes, and opposed
roughened clasping surfaces on the hind limbs.
The common musk turtle is a highly aquatic species and is
found in permanent, and frequently deep, bodies of water. It is
found in many of the clear springs in central Florida, where its
habit of walking on the bottom in search of food can be easily ob
served. This species is very rarely found away from water and is
not normally found on highways even after heavy rain, in contrast
with most species of Kinosternon. However, it does emerge from
the water to bask quite frequently and is often found in precarious
positions on top of cyprees knees or well out along overhanging
branches; from such positions it can fall into the water, when dis
turbed, by simply relaxing its grip. It can remain under water for
long periods, and it is probable that under such circumstances its
highly papillose skin functions as an oxygen exchange surface.
The common musk turtle is almost entirely carnivorous, the
bulk of its diet being made up of carrion, insects, and molluscs.
Small quantities of fish, crayfish, and vegetation may also be
eaten.
561
www.Ebook777.com
Malayan snail-eating turtle, Malayemys subtrijuga. This species reaches a
length of about eight inches. Photo by Dr. Herbert R. Axelrod.
Black pond turtle, Geoclemys hamiltoni, from Southeast Asia. Photograph

ed at the Bronx Zoo.
562
Mediterranean spur-thighed tortoise, Testudo graeca, in its habitat in

Iran.
Large Testudo graeca zarudnyi from central Iran and young Testudo graeca
from North Africa.
graeca
563
www.Ebook777.com
The various vernacular names for the common musk turtle
refer to its habit, when it is picked up or otherwise molested, of
exuding a strong smelling liquid from glands in the soft skin near
the bridge. In my experience, however, the odor is no worse than
that of a number of other turtles, and many specimens seem reluc
tant to emit it at all. After a short period in captivity, most indi
viduals give up the habits of emitting musk and of biting the hand
that holds them.
The common musk turtle hibernates in the more northern
parts of its range; on occasion, many individuals may be found
hibernating together buried in mud under shallow water. Copula
tion may occur in any month during which the turtles are active
and does not appear to be preceded by any elaborate courtship. In
the north, laying is reported to take place from May to August,
while hatchlings appear in September and October. In Florida,’
nesting takes place from April to June and hatchlings are found in
August and September. The eggs are elongate and hard-shelled,
0.95 to 1.22 inches in length and 0.56 to 0.67 inches in width. A clutch
is composed of between one and five eggs. Little effort is made to
bury or conceal the eggs, which are frequently laid between the
buttresses of cypress trees or under logs or dead leaves. Incuba
tion of laboratory-hatched eggs may take from 60 to 75 days.
No subspecies of the common musk turtle have been
described, but Florida specimens tend to be much blacker than
those from further north and they sometimes lose the light lines on
the sides of the head.
The razorback musk turtle, Sternotherus carinatus, is the lar
gest member of the genus, reaching a maximum length of almost
six inches. It is found in streams and swamps throughout Louisi
ana and in adjacent parts of Mississippi, Arkansas, Oklahoma,
and Texas. It has a very elevated, tectiform, flat-sided carapace
with a strong median keel but no lateral keels throughout life. The
carapace is brown, almost uniform in large adults but with radia
ting dark streaks on each scute in young specimens. Some imbri
cation of the scutes is usually apparent. The marginals are low and
are often somewhat eroded in wild specimens. The plastron is
more or less unpigmented in the young, but adults show extensive
darkening of both the plastral scutes and the submarginals. The
gular scute is almost always absent, and there are usually angular
notches at both the anterior and posterior ends of the plastron. The
areas of soft skin between the plastral scutes are especially exten
sive in adult males.
The head is very large, especially in adult males, and is never
striped but has small dark brown spots on the sides and vertical
dark streaks on the jaw surfaces. The snout is strongly projecting.
There is a single pair of barbels near the symphysis of the lower
564
The very high carapace

with a strong median
keel is typical of Ster-
notherus carinatus.
jaw. Differences between the sexes appear to be similar to those

found for S. odoratus.
There appear to be rather few data available on the natural
history of this species. Tinkle found a nest containing two eggs on
the bank of the Peal River, and by dissecting a series of females he
estimated that an average of 7.3 eggs is produced by each female
annually, normally in two clutches. Nesting takes place from April
to June. The hatchling is between 0.9” and 1.2” in length and shows
strongly imbricate carapace scutes and bold spotted patterns on
the head and carapace. Many have traces of lateral keels. The
habitat of the razor-backed musk turtle embraces rivers, slow
streams, and swamps. It prefers situations with little current and
dense aquatic vegetation. Dietarily this species is omnivorous,
feeding mainly on insects and molluscs but also on crustaceans,
carrion, amphibians, and aquatic vegetation. Natural populations
of this species appear to include approximately 60 percent
females. The razor-backed musk turtle, like other Kinosternidae
in the United States, suffers extensive mortality at the hands of
vandalistic fishermen; Mahmoud reported that two fishermen
caught and killed 51 adults of this species in two hours in the Blue
River, Oklahoma.
Two other musk turtles from the southern states, the logger-
head musk turtle and the stripe-necked musk turtle, were former
ly considered subspecies of Stemotherus carinatus. However, al
though they are clearly closely related to each other and appear to
intergrade where their ranges meet, they are now considered to
565
www.Ebook777.com
Hermann’s tortoise, Testudo hermanni, a southern European species fre
quently kept as a pet in Great Britain.
African leopard tortoise, Geochelone pardalis babcocki, from Kenya. This is

one of the largest mainland tortoises. Photo by P. Aldrich-Blake.
566
Radiated tortoise, Geochelone radiata. This tortoise, whose range is

restricted to the island of Madagascar, is the largest and most attractive
of the “starred” tortoises.
Indian starred tortoise, Geochelone elegans. Photographed at the London

Zoo.
567
www.Ebook777.com
belong to a separate species, Stemotherus minor. Intergradation
between minor and carinatus has not been demonstrated, and in
deed the two species appear to be sympatric in southeastern Mis
sissippi, also the gular scute, absent in carinatus, is present in
minor, and the carapaces of the adults of the two species have en
tirely different shapes. The penis structure is also strikingly dif
ferent between the two species.
The loggerhead musk turtle, Stemotherus minor minor, is
found in the northern half of Florida, including all but the tip of the
panhandle, the southern half of Georgia, and southeastern Ala
bama. It is a characteristic and abundant inhabitant of many of
the clear freshwater springs of north-central Florida. This is a
slightly smaller form than the razorback musk turtle, adults being
about four inches long and the record length almost five inches.
Hatchlings have steep, tricarinate shells, light brown in color with
numerous dark brown spots and radiating streaks. As the turtle
grows, the shell becomes relatively lower and more bulbous, but
the three keels remain until old age is approached. The plastron,
which includes a small gular scute, lacks dark pigment but is
bright pink in hatchlings and young, becoming yellow or buff with
growth. The head is grey-brown with numerous small dark spots,
and there are vertical dark streaks on the jaw surfaces. The soft
skin is papillose and maculated, and there are the usual sickle
shaped scales on the front surfaces of the hind limbs and on the
heels. The carapace scutes are slightly imbricate, and the first
central scute is so narrow that it fails by a wide margin to make
contact with the second marginals. The hind part of the tenth, or
last, marginals is considerably raised, and this elevation is con
tinued on the supracaudals.
Differences between the sexes include the much longer
thicker, spur-tipped tail of the male; the male also tends to have a
slightly concave plastron and a head so enlarged that the entire
front part of the shell is distorted to accommodate it.
The loggerhead musk turtle shows an even greater tendency
than the common musk turtle toward basking in almost inacces
sible locations; its small size and reduced plastron allow it to
ascend cypress knees and crawl out along overhanging branches,
where it may remain for hours unless a disturbance prompts it to
relax its grip and drop back into the water.
Experiments by Daniel Belkin of the University of Florida
have revealed that the loggerhead musk turtle is not only capable
of extracting sufficient oxygen for its needs while submerged by
exchanging gases through its thin and papillose skin, but it is also
capable of deriving energy from its food by an anaerobic process
which is more akin biochemically to fermentation than to normal
vertebrate respiration.
568
Nesting of this species apparently takes place in late spring

and early summer. The nests are no more thoroughly constructed
than are those of the common musk turtle. Normal clutches num
ber from one to three eggs.
The diet of the loggerhead musk turtle in the wild has not been
studied systematically, but it is probable that at least the old
males, which have enlarged heads and broad, flat jaw surfaces,
consume considerable quantities of snails and other hard-shelled
invertebrates.
The stripe-necked musk turtle, Stemotherus minor peltifer, is
found in all of Alabama except for the south-east; eastern Missis
sippi; eastern Tennessee; and extreme northwestern Georgia. It is
rather similar to the loggerhead musk turtle but does not reach as
large a size, the record shell length being 4.5 inches. The carapace
of the juvenile is distinctly lower than that of minor, with only
rudimentary lateral keels which disappear much earlier in life.
The carapace of the young is mid-brown with dark spots but no
strong radiating pattern. The head is light yellow-brown with fine
dark spots on the front, but toward the back of the head and on the
neck the pattern becomes one of longitudinal stripes. As with S.m.
minor, the carapace scutes are slightly imbricate, a single pair of
barbels is present on the chin, and a gular scute is present. The
plastron, which is immaculate, has relatively small axillary and
inguinal scutes which are longer than broad. The marking of the
underside of the neck is variable, ranging from distinct longitudi
nal stripes to a reticulate or diffuse pattern of spots.
The stripe-necked musk turtle is highly aquatic, living in
streams and rivers. It has been reported from clear, shallow
creeks up to an altitude of 1000 feet. Its habits in the wild are al
most unknown, but Folkerts (1968) records individuals of peltifer
biting down on mats of algae and riverweed and pulling the algae
back between the jaws to scrape off the tiny snails which covered
the mats of vegetation.
The only remaining member of this genus is a curious, highly
restricted form from northern Alabama. This turtle was described
as Stemotherus depressus by Tinkle and Webb in 1955, but inter
grades with peltifer have now been found and the name has been
altered to Stemotherus minor depressus. The race is only known
from a restricted area of the Black Warrior River system in and
around Walker County, Alabama; the intergrades were found in
the Black Warrior River above Tuscaloosa. It is a small form,
adult at 3 to 4 inches, with so conspicuously flattened and broad
ened a carapace that it looks like a topologically distorted version
of the stripe-necked musk turtle. The carapace has a blunt median
keel and is brown in color with a broken radiating pattern on each
scute in juveniles. The head is finely vermiculated with black on
569
www.Ebook777.com
Red-footed tortoise, Geochelone carbonaria, from northern South America.
Photo by Harald Schultz.
Geochelone denticulata from northwestern Guyana. This is the largest

mainland South American tortoise.
570
Plastral view of Geochelone carbonaria (left) and Geochelone denticulata.
Geochelone carbonaria (left) and Geochelone denticulata.
571
www.Ebook777.com
An inquisitive
depressed musk
turtle, Ster-
notherus minor
depressus.
yellow; the iris of the eye is vivid light yellow and gives the animal
an intense, staring effect. The neck is adorned with dark lines
which vary in number from about 5 to 18. The anterior surfaces of
the forelimbs and the posterior surfaces of the hindlimbs have a
reticulate pattern similar to that of the head. The plastron is ex
tremely small and is attached to the carapace by long narrow
bridges. The plastron is pinkish in young specimens, often with
dark markings (of environmental origin) on the undermarginals.
A single small gular scute is present.
The survival value of the depressed carapace of Sternotherus
minor depressus has not been determined, but it is possible that
the low profile allows the animal to hide in cracks and crevices in
rocks. Suitable rocks for such concealment are abundant in its
habitat, but the 21 specimens representing the type series were
collected in crevices in submerged stumps and in detritus along
the shore.
GENUS STAUROTYPUS
The genus Staurotypus includes the two species of giant musk
turtle from southern Mexico and adjacent parts of Central
America. They are easily distinguished from Kinosternon and
Sternotherus species by the larger size (adults are commonly a
foot in length) and the three strong keels on the carapace. Unlike
all other turtles, the keels on the shells of giant musk turtles be
come stronger with age and correspond to considerable thicken
ings in the bone of the carapace. The shell is also massively con
structed, and it is probable that both the carination and the pachy-
ostosis combine to lessen the likelihood of the turtle being cracked
and eaten by the crocodiles with which it shares much of its habi
tat. Staurotypus has a very large head and a highly reduced plas-
572
tron, and the two species look to the uninitiated somewhat like
snapping turtles. They are, of course, easily distinguished from
true snapping turtles by the narrow, thick shell, the absence of der-
mal papillae, and the presence of a hinge connecting the anterior
part of the plastron to the hyoplastra. Also the axillary and in-
guinal scutes on each side are in contact with a large abdominal
scute which extends as far as the midline of the plastron. The neck
is not particularly long, and there are no free rib-ends.
Staurotypus triporcatus, known in Mexico as 'guau' or 'gala-
pago,' is found in the Gulf and Caribbean drainages of southern
Mexico and northern Central America, having been recorded from
southern Veracruz, Tabasco, Campeche, and northern Chiapas, it
is also known from northern Guatemala and Belize. It is a large
and imposing turtle; fully grown adults of both sexes are about a
foot long. I have seen a fourteen-inch shell from Veracruz, while
the largest on record was a fifteen-inch shell from Roaring Creek,
Belize. Wilfred Neill and Ross Allen, who collected this specimen,
were informed by several local people that one sometimes has to
take to the water after hauling an enraged Staurotypus of large
size into a dugout.
The guau turtle has an elongate, narrow carapace which is
shallow in the young but relatively deep in adults. The three keels
extend the entire length of the shell except for the marginal area;
the median keel in the region of the fourth and fifth centrals is the
highest. There are six neural bones, while the seventh and eighth
pairs of pleurals are completely separated along the midlme by an
elongate suprapygal. The carapace is brown, with dark blotches
Juvenile guau turtle, Staurotypus triporcatus, from the Caribbean drainages

of Central America.
573
www.Ebook777.com
Argentine tortoise, Geochelone chilensis. This is the smallest of the three
tortoise species found in mainland South America. Photo by Dr. C. M. Bogert.
Galapagos tortoise, Geochelone elephantopus elephantopus, from South

Albemarle Island. Photographed at the Bermuda Aquarium.
574
Female Galapagos tortoise, Geochelone elephantopus becki, from North

Albemarle Island. Males of this race have developed a strongly saddle-
shaped carapace. Photographed at the Bronx Zoo.
575
www.Ebook777.com
forming a vague radiating pattern on each scute, concentrated
toward the areola. The marginal scutes are very low anteriorly but
become progressively higher toward the rear. The yellow plastron
is composed of only four pair of scutes, the second pair having ex
tensions that make up the inner part of the bridge. The axillary
and inguinal scutes are large and have a considerable line of con
tact. The rounded front plastral lobe is attached by a hinge which
forms a strongly sinuous line in specimens more than about six
inches in length. The back of the plastron terminates in a sharp
point. In some specimens there may be traces of dark markings on
the plastral scutes.
The head is very large and appears to become progressively
wider with age; a twelve-inch specimen has a head about 3*4
inches wide. The snout is very pointed, and its upper surface is pro
tected by a single scale which extends backward over each eye.
The jaws have strong crushing surfaces and are totally lacking in
ridges and denticulations. The head is black, with numerous prom
inent white spots; the jaws are either uniformly black or are
heavily blotched with black streaks. There is a pair of barbels on
the chin.
Adult Staurotypus
triporcatus have a
distinct snout.
The limbs are of medium size, and the toes are fully webbed.
There is a series of three or four curved, sickle-like scales of de
creasing size on the lower foreface of each front limb and more
numerous scales of similar shape on each heel. Males have
opposed patches of tubercular scales on the tibial and femoral
parts of the hind limbs, as well as long, thick-based tails with the
vent relatively distally placed. Males also have narrower bridges
than the females. The soft parts of the guau turtle are dark grey
dorsally, creamy-yellow or white ventrally.
576
The guau turtle lives in large lakes and lagooons; Guatemalan

specimens are often covered with a thick growth of algae, but this
seems to be rare in Veracruz. This species when newly captured
often excretes sizeable portions of the shells of the smaller mud
turtles with which it shares its range—Kinosternon leucostomum,
acutum, etc.; its massive jaws enable it to crack up small turtles
without difficulty. It also feeds upon heavy-shelled molluscs of
various sorts. It in turn is liable to be eaten by Morelet’s crocodile
(Crocodylus moreleti) which, at least in former times, was found
throughout the range of the guau. It is probable that the thick shell
and strong keels are a response to this predation. The meat of this
turtle is also greatly appreciated for human consumption in all
parts of its range.
The eggs of the guau are hard-shelled and elongate, about 1.5”
in length and 0.9” in width. There are relatively few in a clutch—
probably usually between three and six. The hatchlings are about
1.2 inches in carapace length and have large black blotches on the
carapace, submarginals, bridge, and plastron. The species has
bred in captivity at Ross Allen’s Reptile Institute.
The crucilla turtle, Staurotypus salvini, is the Pacific coast
relative of the guau; it is found in the coastal lowlands of the states
of Oaxaca and Chiapas, as well as in Guatemala and El Salvador.
Most of the published keys to differentiate the two species rely on
characteristics of the relative proportions of parts of the plastron,
which in fact vary with both age and sex. For example, the width
of the bridge has been used to differentiate the species, salvini sup
posedly having the narrower bridge, but it appears likely that the
originator of this distinction simply compared a male salvini with
a female triporcatus. It has also been reported that triporcatus has
a strongly sinuous plastral hinge while salvini has a relatively
straight one, but here again a series of triporcatus shows increas
ing sinuosity with age. The relative proportions of the abdominal
scutes and the particular pair of scutes which shows the longest
median suture have also been used, unsuccessfully, to separate the
species.
The two species of Staurotypus are in fact best distinguished
on the basis of the coloration of the head. S. triporcatus has black,
or almost black, jaw surfaces and prominent white spots on the
head. S. salvini has fine orange-yellow spots on the head, light yel
low jaw surfaces, and a light area in the soft skin near the comer of
the mouth. S. salvini has a more aggressive disposition than tripor
catus and shows an even greater inclination to bite the hand that
holds it.
Boulenger (1889) wrote that the seventh pleural bones of sal
vini have a median suture behind the seventh neural bone, and this
situation is illustrated (p. 31). However, this must be a rare vari
ant condition; those specimens of salvini that I have examined had
577
www.Ebook777.com
Geochelone elephantopus chathamensis from Chatham Island in the
Galapagos. Photographed at the Bronx Zoo.
Aldabra Island giant tortoise, Geochelone gigantea. This species is plentiful

on Aldabra in the Indian Ocean. Photographed at Ross Allen’s Reptile In
stitute.
578
Saddle tortoise, Geochelone elephantopus ephippium, from Duncan Island in

the Galapagos. Photographed at the Bronx Zoo.
Indefatigable Island tortoise, Geochelone elephantopus porteri. This is the

most abundant of the surviving races of Galapagos tortoises. Photographed
at the St. Augustine Alligator Farm.
579
www.Ebook777.com
exactly the same carapace bone configuration as is typical of tri-
porcatus.
I have never seen a salvini as large as a large triporcatus, and
it is likely that the adult size is substantially smaller. The habits of
the crucilla turtle have not been studied, but the species in cap
tivity will eat virtually any animal matter offered. The species has
bred in captivity in the collection of Alfred A. Schmidt in Ger
many. Schmidt's breeding female was 7.9 inches in length and the
male 7.2 inches—smaller than the normal breeding size of tripor
catus. The pair was seen copulating, in water, on January 14, 1968;
because of her relatively deep shell and posterior carapace over
hang, the male was unable to reach the front margin of the
female’s carapace with his front limbs; however, he maintained a
firm grip on the back of the female’s shell with his hind limbs, the
opposing patches of homy tubercles providing adequate friction
for a good grip. During the performance the female reached up
ward and gently bit the fully extended head of the male. Nine eggs,
each measuring approximately 1.6 by 0.8 inches, were laid on Sep
tember 18, and six more on December 27. Of these, three hatched
after about 207 days incubation. The hatchlings weighed between 5
and 7 grams and had carapace lengths ranging from 1 inch to 1.2
inches. Within a month the two hatchlings that survived doubled
their hatchling weights and increased about 0.4 inches in carapace
length.
GENUS CLAUDIUS
The narrow-bridged mud turtle, Claudius angustatus, is a
strange little animal that would appear to be most closely related
to Staurotypus, but which nevertheless shows several striking and
unusual features. This species has a rather flat, oval shell, very
variable in relative width and either parallel sided or wider pos
teriorly than anteriorly. The scutes of the carapace are drab yel
low-olive, but each is adorned with a pattern of radiating black
spots and broken streaks. This pattern is not usually present in
those areas at the edges of the scutes corresponding to the most
recent growth, and in some specimens the pattern may become so
obscure that it almost disappears. The marginal scutes form an
exceedingly low rim around the edge of the shell, and as with all
mud turtles there are only ten marginal scutes on each side (not
counting the paired supracaudals). The shell bears traces of three
longitudinal keels even in large specimens, but these are so feeble
that, unlike Staurotypus, they can in no way contribute to the
mechanical strength of the shell. The neural bones number six or
seven; the first one is narrow anteriorly and only just makes con
tact with the nuchal bone. The second neural is large and often
octagonal, though in other specimens it may be heptagonal or hex-
580
Claudius angustatus has a unique reduced plastron with only seven bones.
581
www.Ebook777.com
Geochelone emys from Thailand and Burma. Drawing by Peter Parks.
West African hinged tortoise, Kinixys belliana. The genus Kinixys is the only
one whose members have a hinge in the caparace. Photo by Dr. Herbert R.
Axelrod.
582
Elongated tortoise, Geochelone elongata, from India.
Travancore tortoise, Geochelone travancorica. This species is very similar to

Geochelone elongata but lacks the nuchal scute.
583
www.Ebook777.com
agonal. The shape of the third neural depends upon that of the
second, but the neurals toward the rear of this are typically hex
agonal with the short sides in front, the last of all being pentagonal.
The sixth pair of pleurals sometimes makes a median suture, the
seventh pleurals always do, and the eighth sometimes do, while in
other cases they are separated mesially by extensions of the
seventh pleurals which extend back to meet the suprapygal.
The plastron of Claudius is highly reduced and is unique in
being composed of only seven bones; a well-developed entoplas-
tron and paired entoplastra and xiphiplastra are present, but the
area occupied by the hyoplastra and hypoplastra of other turtles is
occupied by a single pair of bones. The lateral borders of these
bones are extended outward and backward into the long and ex
ceedingly narrow bridges, which form no bony suture with any
part of the carapace but instead are attached to the carapace by
flexible ligamentous material. The outer ends of the bridges are
equipped with strong backwardly directed processes to extend the
line of contact of the bone with the cartilage. The plastron is
covered with only four pairs of scutes, the bridges being partially
covered by extensions from the posterior corners of the second
pair. The outer parts of the bridges are covered with soft skin, and
no axillary and inguinal scutes are present; there may be rudi
mentary vestiges of a minute gular scute at the front of the plas
tron. The plastron is yellow, though in some individuals it may be
stained dark reddish by ferric deposits. Both carapace and plas
tron of Claudius are very thin, and though the bridges are reason
ably stout, their flexible connection with the carapace gives the
impression that the turtle would be very easy to crush with only
moderate pressure.
The head of Claudius is very large, about half the width of the
carapace. The skull is elongate, with the eyes well forward and the
snout pointed. The upper jaw has a small median point and
another tooth-like pointed projection on each side just below the
front corner of the eye—a feature unique among turtles. The lower
jaw has an extraordinarily long median hook—so long that the
cavity in the upper jaw which accommodates this hook when the
mouth is closed extends right into the nasal chamber. This in turn
causes the upper surface of the tip of the snout to be slightly bul
bous, and the swelling is protected externally by a single oval
scale. The tongue is thick and fleshy and is attached throughout its
length to the floor of the mouth. There is a pair of small barbels on
the chin. The feet are small but the toes are fully webbed. Apart
from the rhamphothecae and the scale on the top of the snout, the
skin of Claudius is virtually scaleless except for three wide,
curved, band-like scales on the lower part of the front face of the
forelimbs near the digits and a patch of similar scales in the heel
584
The strongly cusped jaws of Claudius angustatus are one hint that it might
feed on frogs in nature.
area. Also, males have opposed patches of small, rough scales on

the rear surfaces of the femoral and tibial portions of the hind
limbs. The males also differ from the females in having long pre
hensile tails with thick bases and hard, homy tips. There does not
appear to be any significant difference between the sizes reached
by the two sexes, though unusually large specimens seem to be
mostly males; adults of both are typically between four and five
inches long, while really large individuals may be 5.5 to 6 inches in
length.
The coloration of the soft parts of Claudius is typically dark
grey-black dorsally and light grey to buff or cream ventrally. The
jaw plates are yellow with vertical dark streaks, and there may be
light spotting or vermiculation on the comers of the mouth and the
chin and on the scale on top of the snout.
Claudius has been recorded from the Mexican states of Vera
cruz (not north of the City of Veracruz), Tabasco, and Campeche,
585
www.Ebook777.com
Mexican gopher tortoise, Gopherus flavomarginatus, a recently discovered
species.
Gopher tortoise, Gopherus polyphemus, a burrowing species found in sandy

areas of the southeastern United States. Photographed at Ross Allen’s Rep
tile Institute.
586
Homopus areolatus, a
small South African tor
toise. Photo by Mervin
F. Roberts.
Desert tortoise,
Gopherus agassizi,
found in arid areas of the
southwestern United
States. Photo by
Dr. C.M. Bogert.
Berlandier’s tortoise,
Gopherus berlandieri, a
small species that is
very abundant in Texas.
www.Ebook777.com
as well as from Belize. The species is presumably also found in
Guatemala and possibly also in Quintana Roo. In Mexico, it is
known as ‘Talmama’ or ‘Chopontil;’ despite its very small size, it
is used for human consumption.
The life history of Claudius is very poorly known; this is a pity
since several of its features suggest habitat and diet specializa
tion. It is known that the breeding season coincides with the onset
of the dry season in November; Peter Hausmann found that fe
males from an undisclosed part of Mexico laid eggs soon after cap
ture in November, and he ascertained that between two and eight
very small ‘oblong-shaped’ eggs were deposited at a time. A
female Claudius with a carapace about 4.75 inches in length that I
purchased in Veracruz at the end of December was found after
death to contain three elongate hard-shelled eggs about 1.25” in
length and about 0.75” wide, as well as five ovarian yolks each
about 0.5” in diameter. Another female with a 4.63 inch carapace
contained a single shelled egg and ten ovarian yolks ranging stead
ily in size from the smallest, about 0.3” in diameter, to the three
largest, each about 0.75” in diameter. From these data it would
appear that Claudius nests more than once in a season. Hausmann
found that the eggs took five months to hatch at 28° C and that the
hatchlings initially had black dorsal surfaces which became brown
after a few days.
Hausmann’s specimens were found in a marshy area formed
by a slow-moving river, and it may be that such a coastal flood-
plain type habitat is typical of the species throughout its range;
this might explain the lack of definite locality records from
northern Guatemala. William Rainey informs me that he found
Claudius in a small rather steep-banked stream close to the city of
Belize. Claudius, as mentioned above, has such a thin shell and
such exposed soft parts that it would tend to fall easy prey to croco
diles if it shared their habitat. Those turtles which do co-exist with
Morelet’s crocodile in this area (Staurotypus triporcatus, Pseu-
demys scripta venusta, Kinosternon integrum) have very strong,
thick shells. Possibly Claudius, by living in marshes, shallow pud
dles, streams, etc., avoids coming in contact with crocodiles,
which tend to live in deeper water.
Captive Claudius feed quite readily on fish, beef, shrimps,
worms, etc., but no data are available on the diet of the species in
the wild. However, certain morphological and behavioral speciali
zations of the animal allow us to make some speculations in this
connection. Claudius has the barbels on the chin which one asso
ciates with the bottom-walking, predatory approach to life in tur
tles. Moreover it has an extremely long neck with the well-devel
oped longissimus dorsi muscles and free rib-ends within the shell
which allow it to make a snapping-turtle-like strike. The mouth is
588
very large, but the jaw surfaces with their array of tooth-like
spikes and sharp edges are uniquely adapted for holding and cut
ting, rather than crushing prey. It would appear that Claudius
feeds upon highly active but rather soft-bodied animals. In view of
the marshy habitat of the species, it appears not unlikely that it
may eat frogs in some numbers.
Elizabeth Wing informs me that Claudius in Veracruz may
bury itself in the mud when the marshes in which it lives dry up in
November and may remain buried for several months. Local
people locate buried specimens by probing with rods; apparently
they do not see the species at all during the wet season, when the
marshes are flooded to depths of several feet.
Claudius is delicate in captivity, though occasionally one will
live for several years. Captive specimens must be handled with
care, since the neck is so long that the turtle can bite almost as far
back as its own tail.
BACKGROUND READING
CHELYDRIDAE AND KINOSTERNIDAE (U.S. SPECIES):
Carr, A.F. 1952. Handbook of Turtles. Comstock Publ. Assoc.,
Ithaca, N.Y. 542 pp.
States. Kentucky Univ. Press, Lexington. 347 pp.
Pope, C.H. 1939. Turtles of the United States and Canada.
Alfred A. Knopf, Inc. New York. 343 pp.
MEXICAN SPECIES:
Casas-Andreu, G. 1967. Contribucion al conocimiento de la tor-
tugas dulceacuicolas de Mexico. Tesis professional, Univ.
Nac. Mexico. 96 pp.
Smith, H.M. and E.H. Taylor. 1950. An annotated checklist and
key to the reptiles of Mexico exclusive of the snakes. Bull.
U.S. Nat. Mus. No. 199, i-v, 1-253.
CHELYDRA SERPENTINA:
Alvarez del Toro, M. 1960. Los Reptiles de Chiapas. Tuxtla
Gutierrez, Chiapas, Mexico. 204 pp.
Feuer, R.C. 1966. Variation in snapping turtles, Chelydra ser
pentina Linnaeus: a study in quantitative systematics.
Ph.D. thesis, University of Utah, Salt Lake City.
__________. 1971. Intergradation of the snapping turtles Chely
dra serpentina serpentina (Linnaeus, 1758) and Chelydra
serpentina osceola Stejneger, 1918. Herpetologica 27: 379-
384.
589
www.Ebook777.com
Swimming male green turtle, Chelonia mydas. Note the streamlined form and
very long tail. Photographed at Marineland of Florida.
Hatchling loggerheard turtle, Caretta caretta. The species still nests in

numbers on the coasts of Georgia and Florida. Photo by D. Faulkner.
590
Atlantic loggerhead turtle, Caretta caretta, easily identified by the large head
and the reddish brown coloration. Photographed at Marineland of Florida.
An adult female green turtle, Chelonia mydas, returning to the sea after
nesting the night before on Shell Beach, Guyana.
www.Ebook777.com
Medem, F. 1962. Le distribucion geographica y ecologia de Los
Crocodylia y Testudinata en el departamento del Choco.
Rev. Acad. Colombiana Ciencias Exactas, Fisicas y Nat
11(44): 279-303.
Richmond, N.D. 1958. The status of the Florida snapping turtle
Chelydra serpentina osceola Stejneger. Copeia 1958: 41-43
MACROCLEMYS TEMMINCKII:
Allen, E.R. and W.T. Neill. 1950. The alligator snapping turtle,
Macrochelys temminckii, in Florida. Spl. Publ. Ross Allen’s
Rept. Inst. 4: 1-15.
Dobie, J.L. 1971. Reproduction and growth in the alligator snap
ping turtle, Macroclemys temminckii (Troost). Copeia (4V
645-658.
Hall, H.H. and H.M. Smith. 1947. Selected records of reptiles
and amphibians from southeastern Kansas. Trans. Kansas
Acad. Sci., 49: 447-454.
Minton, S.A. and M.R. Minton. 1973. Giant reptiles. Charles
Scribner’s sons, New York, 345 pp.
nois Natural History Survey Bulletin 28: 1-298.
Wickham, M.M. 1922. Notes on the migration of Macrochelys
lacertina. Oklahoma Acad. Sci., n.s., 247, Univ. Studies 15-
20-22. ’ '
KINOSTERNON SUBRUBRUM:
Mahmoud, I.Y. 1969. Comparative ecology of the kinostemld
turtles of Oklahoma. S.W. Nat. 14: 31-66
KINOSTERNON BA URI:
Einem, G.E. 1956. Certain aspects of the natural history of the
mud turtle Kinosternon bauri. Copeia 1956: 186-188.
Uzzell, T.M. and A. Schwartz. 1955. The status of the turtle
Kinosternon bauri palmarum Stejneger with notes on varia
tion in the species. J. Elisha Mitch. Soc. 71: 28-35
KINOSTERNON FLAVESCENS:
Hartweg, N. 1938. Kinosternon flavescens stejnegeri, a new tur
tle from northern Mexico. Occ. Papers Mus. Zool. Univ
Michigan 371: 1-5.
Mahmoud, I.Y. 1969. op. cit.
Punzo, F. 1974. A qualitative and quantitative study of the food
items of the yellow mud turtle, Kinosternon flavescens
(Agassiz). Journal of Herpetology 8(3): 269-270.
KINOSTERNON SONORIENSE:
Hulse, A.C. 1974. Food habits and feeding behavior in Kinoster
non sonoriense (Chelonia: Kinosternidae). Journal of Herp
etology 8(3): 195-200.
592
__________in press. The aquatic turtles of Arizona.

Niles, D.M. 1962. Records for the Sonora mud turtle, Kinoster
non sonoriense, in New Mexico. Herpetologica 18: 205-206.
Van Denburgh, J. 1922. The reptiles of western North America.
Vol. II. Snakes and Turtles. Occ. Papers Calif. Acad. Sci.,
10: 623-1028.
KINOSTERNON LEUCOSTOMUM:
Medem, F. 1962. op. cit.
Moll, E.O. and J.M. Legler. 1971. The life history of a neotropi
cal slider turtle Pseudemys scripta (Schoepff) in Panama.
Bull. Los Angeles County Mus. Nat. Hist. Sci.: No. 11: 1-102.
KINOSTERNON SCORPIOIDES:
Alvarez del Toro, M. 1960. op. cit.
da Cunha, O.R. 1970. Uma nova subespecie de quelonio, Kino
sternon scorpioides carajasensis de Serra dos Carajas,
Para. Bol. Mus. Paraense Emilio Goeldi, n.s. Zoologica,
No. 73: 1-12.
Freiberg, A. 1936. Una nueva tortuga del norte argentmo. Phy-
sls, Buenos Aires, 41: 169-171.
Muller, L. and W. Hellmich. 1936. Amphibien und Reptilien, 1.
Teil: Amphibia, Chelonia, Loricata. Wiss. Ergebnlsse
Deutsche. Gran Chaco-Exped. Stuttgart. 120 pp.
Sexton, O.J. 1960. Notas sobre la reproduccion de una tortuga
Venezolana, la Kinosternon scorpioides. Mem. Soc. Cien.
Natur. 20(57): 189-197.
KINOSTERNON INTEGRUM:
Casas-Andreu, G. 1967. op. cit.
LeConte, J. 1854. Descriptions of four new species of Kinoster-
num. Proc. Acad. Nat. Sci. Philadelphia. Vol. VII: pp. 180-
190.
KINOSTERNON ACUTUM:
Neill, W.T. 1965. New and noteworthy amphibians and reptiles
from British Honduras. Bull. Florida State Mus., 9, No. 3.
77-130.
KINOSTERNON HIRTIPES:
Duellman, W.E. 1965. A biogeographic account of the herpeto-
fauna of Michoacan, Mexico. Univ. Kansas Publ. Mus. Nat.
Hist. 15: 627-709.
Glass, B.P. andN. Hartweg. 1951. Kinosternon murrayi, a new
muskturtle of the hirtipes group from Texas. Copeia 1951:
50-52.
KINOSTERNON ABAXILLARE:
Stejneger, L. 1925. New species and subspecies of American
turtles. J. Washington Acad. Sci., 15: 462-463.
Alvarez del Toro, M. 1960. op. cit.
593
www.Ebook777.com
Head of a leatherback turtle, Dermochelys coriacea, from Bigisanti, Surinam.
The leatherback is the largest living turtle.
Ridley sea turtle, Lepidochelys olivacea, from Shell Beach, Guyana.

Atlantic ridley, Lepidochelys kempi, from the Gulf of Mexico. This ridley,
smallest of the sea turtles, is known for its tremendously aggregated nesting
habits. Photographed at Marineland of Florida.
Half-grown hawksbill turtle, Eretmochelys imbricata, from Grand Cayman

Island.
595
www.Ebook777.com
KINOSTERNON CREASERI:
Duellman, W.E. 1965. Amphibians and Reptiles from the Yuca
tan Peninsula, Mexico. Univ. Kansas Publ. Mus. Nat. Hist.
15, No. 12: 577-614.
Hartweg, N. 1934. A new mud turtle from Chichen Itza, Yuca
tan. Occ. Pap. Mus. Zool. Univ. Mich., Ann Arbor. No. 277: 1.
KINOSTERNON HERRERAI:
Pritchard, P.C.H. 1969. Herrera’s mud turtle. Int. Turt. Tort.
Soc. J. 3, No. 3: 6-9, 35.
Stejneger, L. 1925. op. cit.
KINOSTERNON ANGUSTIPONS:
Legler, J.M. 1965. A new species of turtle, genus Kinosternon,
from Central America. Univ. Kansas Publ., Mus. Nat. Hist.
15, No. 13, 615-625.
__________ 1966. Notes on the natural history of the rare Central
American turtle Kinosternon angustipons Legler. Herpeto-
logica, 22, No. 2: 118-122.
KINOSTERNON DUNNI:
Medem, F. 1961. Contribuciones al conocimiento sobre la mor-
fologia, ecologia, y distribucion de la tortuga Kinosternon
dunni K.P. Schmidt. Novedades Colombianas, 1(6): 446-476.
__________ 1962. op. cit.
Schmidt, K.P. 1947. A new kinostemid turtle from Colombia.
Fieldiana Zoology, 31(13): 109-112, fig. 14.
STERNOTHERUS ODORATUS:
Risley, P.L. 1933. Observations on the natural history of the
common musk turtle, Stemotherus odoratus (Latreille).
Pap. Michigan Acad. Sci. Arts Lett. 17: 685-711.
Tinkle, D.W. 1961. Geographic variation in reproduction, size,
sex ratio, and maturity of Stemotherus odoratus (Testudi-
nata: Chelydridae). Tulane Stud. Zool. 9: 331-349.
STERNOTHERUS CARINATUS:
Tinkle, D.W. 1958. The systematics and ecology of the Stemo
therus carinatus complex (Testudinata: Chelydridae).
Tulane Stud. Zool. 6: 3-56.
STERNOTHERUS MINOR:
Belkin, D.A. 1968. Aquatic respiration and underwater survival
of two freshwater turtle species. Resp. Physiol. 4: 1-14.
Folkerts, G.W. 1968. Food habits of the stripe-necked musk tur
tle, Stemotherus minor peltifer Smith and Glass. J. Herpe-
tol. 2: 171-173.
Tinkle, D.W. and R.G. Webb. 1955. A new species of Stemo
therus with a discussion of the Stemotherus carinatus com
plex (Chelonia. Kinosternidae). Tulane Stud. Zool. 3: 53-67.
596
STAUROTYPUS TRIPORCATUS:
Neill, W.T. and E.R. Allen. 1959. Studies on the amphibians and
reptiles of British Honduras. Publ. Research Div., Ross
Allen’s Reptile Inst., vol. 2, no. 1, pp. 1-76.
Neill, W.T. 1965. op. cit.
STAUROTYPUS SALVINI:
cephalians, and crocodiles in the British Museum. Pp. iii-x,
1-311.
Schmidt, A.A. 1970. Zur Fortpflanzung der Kreuzbrustschild-
krote (Staurotypus salvini) in Gefangenschaft. Salamandra,
Band 6, Heft 1/2, pp. 3-10.
Stuart, L.C. 1963. A checklist of the herpetofauna of Guatemala.
Misc. Publ. Mus. Zool. Univ. Mich., no. 122, pp. 1-150.
CLAUDIUS ANGUSTATUS:
Hausmann, P. 1968. Claudius angustatus. Int. Turt. Tort. Soc.
J. 2, No. 1: 14-15.
597
www.Ebook777.com
Large adult Florida soft-shelled turtle, Trionyx ferox. Photographed at Ross
Allen’s Reptile Institute.
Gulf Coast soft-shelled turtle, Trionyx spinifer asper. Photographed at the

London Zoo.
598
Ganges soft-shelled turtle, Trionyx gangeticus. Adults are esteemed as food.
Nile soft-shelled turtle, Trionyx triunguis. Photo by S. Mathiasson.
599
www.Ebook777.com
Chapter 7
SOFT-SHELLED TURTLES
The soft-shelled turtles form the family Trionychidae, which is
not named for the scuteless, flexible carapace, but for the much
less conspicuous feature of the presence of only three claws on
each foot. The distribution of the family is roughly similar to that
of the Emydidae; soft-shells are abundant in temperate eastern
North America and tropical Southeast Asia. They are also found on
the more western islands of Indonesia, one species is found in the
Middle East, and there are several in Africa. Soft-shells are not
even known as fossils from any part of the Australian continent,
and until recently this was also thought to be true of South Ameri
ca. However, my friend Roger Wood recently showed me a well-
preserved fossil hypoplastron of a large Trionyx which he had
found in Venezuela. The group is a very ancient one, despite its
very advanced features, and was already extant in the late Juras
sic.
Soft-shells are often large—sometimes very large—turtles
whose general appearance has often been compared to that of an
animated pancake. This comparison is not unreasonable, since
most soft-shells have scuteless, discoidal, flat, flexible "shells;”
but it should be remembered that only the rim of the carapace is
flexible, the entire antero-central part being supported by con
tinuous—and often thick—bone, while the sides are strengthened
by the ends of the flattened ribs. Also, some soft-shells, such as the
Florida soft-shell, do not have particularly broad shells, while
others (such as the Indian flap-shell, Lissemys punctata) are quite
strikingly dome-shaped. Pathological formation of an unusually
domed carapace (kyphosis) is also more common among other
soft-shell species than among hard-shelled turtles.
Soft-shells are fully aquatic animals, generally very wary in
the wild, and capable of rapid locomotion both on land and in
water. They have soft skin with scales reduced to a few sickle
shaped lamellae on the limbs, and irregularities and projections
on the carapace of some species that may or may not be homolo
gous to true scales. Peripheral bones are completely absent, ex
cept for rudiments in the single species Lissemys punctata, and
the pleural bones are usually reduced to seven pairs in the four
600
New World species. The neural bones show only slight reduction,
the posterior pair of pleurals only usually showing a median line of
contact. The hones of the plastron in young soft-shells are reduced
to splint-like rudiments that cover only a fraction of the total plas-
tral area; even in adults extensive boneless areas remain. The
entoplastron is absent, the epiplastra are fused into a boomerang
shaped bone, and there is a pair of neomorphic bones, known as
‘preplastra,’ in front of the fused epiplastra. The hyo- and hypo-
plastra have a very close sutural connection, and these bones may
become completely fused in old adults of some species. The front
part of the plastron of soft-shells may be raised quite as effectively
601
www.Ebook777.com
Chinese soft-shelled turtle, Trionyx sinensis. Photo by Dr. Herbert R. Axelrod.
Indian soft-shelled turtle, Lissemys punctata. The shell is unusually highly

domed for a soft-shell, and the plastron has fleshy flaps to protect the limbs.
Photo by Dr. Herbert R. Axelrod.
602
Adult Podocnemis unifilis from Guyana. Photo by Dr. Herbert R. Axelrod.
Young yellow-spotted Amazon turtle, Podocnemis unifilis. Photo by Dr.

Herbert R. Axelrod.
www.Ebook777.com
as that of hinged hard-shell species such as Pyxis, but because of
the generally reduced bony structure of the plastron, the line of
flexion is not obvious in prepared skeletons.
Soft-shells have long, highly mobile and flexible necks; they
are also frequently aggressive in disposition, so holding a soft-
shell, a difficult task at best in view of the slippery nature and
active struggles of the beast, is also somewhat dangerous since the
animal can reach amazingly far back to bite its holder. The neck
vertebrae are all opisthocoelous except for the eighth, which is
convex in front and has a ligamentous non-articular connection
with the first trunk vertebra behind. Such a connection is neces
sary, since in full retraction the haemal surfaces of the eighth cer
vical and the first dorsal vertebra are adpressed. The joint behind
the seventh cervical is always doubled, that between the seventh
and sixth sometimes so. The soft-shell skull is elongated, strongly
emarginated in the temporal region, and with rather dorsally-
located orbits. The jaw surfaces may be equipped either with
sharp cutting edges or with flattened grinding surfaces; in either
case, the true nature of the jaws is concealed in living animals by
soft, fleshy lips. The snout ends in a proboscis that may be long and
thin or relatively short and stout; such a proboscis is otherwise
found among turtles only in Chelus and Carettochelys.
Soft-shells lack cloacal bursae and hence do not indulge in anal
respiration. They definitely take in water through the mouth and
expel it through the nose, but this may be at least partially for ol
factory as opposed to respiratory purposes. It is possible that soft-
shells conduct a fair amount of underwater respiration through the
skin; the soft, delicate skin would seem to be somewhat of a lia
bility otherwise, and the soft, unpigmented undersurfaces of living
soft-shells often show extensive capillary networks very close to
the surface.
Subadult and adult soft-shells show strongly sculptured sur
face bones, or epithecal ossifications, in the shell. Those of the
carapace correspond closely with the normal ‘thecal” shell bones
(the neurals, pleurals, etc.) but those of the plastron may differ
markedly in both number and shape from the thecal bones of the
plastron.
The relationships of the Trionychidae are not immediately
obvious. It appears not unlikely that the family is related to the
Carettochelyidae, whose sole living representative is the plateless
river turtle from New Guinea. The families differ, however, in that
Carettochelys has an essentially normal bony shell structure with
a completely ossified rim composed of well-developed peripheral
bones, and a large plastron, lacking fontanelles, and with an ento-
plastron, separate epiplastra, and no preplastra. On the other
hand, Carettochelys resembles the soft-shells in having a layer of
604
soft skin covering the entire shell instead of the usual horny scutes.
Moreover, the shell bones of Carettochelys are strongly sculptured
on their outer surfaces and the snout terminates in a blunt but dis
tinct ‘snorkel.’ The soft-shells are also considered by some authors
to show relationships with the Dermatemyidae and the Chelydri-
dae.
The six genera of living soft-shells may quite reasonably be
separated into two subfamilies: the Cyclanorbinae, including the
African Cyclanorbis and Cycloderma and the Indian Lissemys,
and the Trionychinae, containing the widespread Trionyx and the
Southeast Asian Pelochelys and Chitra. The two groups are distin
guished by the presence in the former only of fleshy femoral flaps
which can be closed to conceal the hind limbs. The survival value
afforded by *hese flaps is obscure, since they are so soft that they
would seem to offer no protection against the attacks of a deter
mined predator.
Lissemys is distinguished from other members of its sub
family by the presence of small isolated bones in the margin of the
carapace; these are very probably homologous to the peripheral
bones of hard-shelled turtles. In this genus seven plastral callosi
ties are normally present, but the two xiphiplastral callosities may
fuse mesially, reducing the total to six. Cyclanorbis lacks the peri
pheral bones, has a postorbital arch narrower than the diameter of
the orbit, long and angular preplastra, and seven plastral callosi
ties in adults.
The genera of the Trionychinae are distinguished by the posi
tion of the orbits; in Trionyx the postorbital arch is narrower than
the diameter of the orbit, in Pelochelys the postorbital arch is
slightly broader than the orbit, and in Chitra the orbits are located
at the very front of the strongly elongated skull, so that the post
orbital bar is several times the width of the orbit. The invalid
genus Dogania, which has now been subsumed within Trionyx, has
an extremely narrow postorbital bar, but this condition is
approached by several other species of Trionyx (e.g. muticus).
Soft-shells of the genus Trionyx are even today often referred
to by the generic name Amyda. The two names were proposed by
different authors, Geoffroy and Schweigger respectively, in the
same year (1809). The name Trionyx antedates Amyda by a few
months, but there is some argument as to which species should be
considered the type. Some maintain that the type was designated
as Trionyx aegyptiacus ( = T. triunguis), but others insist that the
type should be T. coromandelicus. The problem is that the former
of these species is a true Trionyx, but the latter is now considered a
synonym of Lissemys punctata granosa. Consequently, those who
consider coromandelicus the type species of Trionyx insist that the
genus normally known as Lissemys should be called Trionyx, and
605
www.Ebook777.com
Young Podocnemis erythrocephala from South America. Adults lack the red
markings on the head. Photo by Dr. Herbert R. Axelrod.
Young arrau river turtle, Podocnemis expansa, from Guyana. Photo by Dr.
Herbert R. Axelrod.
606
Podocnemis lewyana, from Venezuela, is a rare species lacking the promi

nent head spots of its more familiar relatives.
African mud turtle, Pelusios subniger. Photo by Dr. Herbert R. Axelrod.
607
www.Ebook777.com
that which Is normally called Trionyx should be referred to by the
next available name, Amyda. The choice of names used in this
book reflects the majority opinion today.
GENUS LISSEMYS
The Indian flap-shelled turtle, Lissemys punctata, exists as
three subspecies. L.p. punctata is known from the Indus and
Ganges River Systems, from the Northwest Frontier Provinces
east to Sikkim and south to Kutch and eastern Bangladesh. L.p.
granosa inhabits peninsular India and Ceylon, while L.p. scutata is
restricted to Burma and Pegu. Lissemys punctata is one of the
smallest of the soft-shells; males are mature at less than six
inches, while the largest females are eleven inches long and weigh
about ten pounds. This species has a rather domed shell which is
virtually unflared and has very little of the flexible flap so typical
of soft-shells. The carapace of L.p. punctata is of a grey-green
color with numerous black-bordered yellow spots of irregular
arrangement and size. The first marginal bone is much larger
than the second, and there is a prenuchal bone and a small ento-
plastral callosity. The carapace has a light rim and the underside
is unpigmented. The head and snout are rather short, with a well-
defined, mesially broken upper lip. The head is grey-green with
prominent yellow blotches and streaks on the sides. The surface of
the carapace is tuberculate in young specimens, the tubercles
being roughly aligned to form broken longitudinal ridges. As with
The yellow spotting makes the Indian Lissemys punctata punctata the most
attractive of the flap-shelled soft-shells.
Two views of Lissemys punctata scutata. The plastral callosities are clearly
shown in the photo above. Specimen from the Lantz collection.
www.Ebook777.com
The turtles of the genus Pelusios have a hinged plastron. Photographed at
the London Zoo.
African side-necked turtle, Pelusios adansoni. Photographed at the London

Zoo.
610
An Australian side-necked turtle, Emydura kreffti. Photo by Dr. C. M. Bogert.
Australian mollusc-eating turtle, Emydura australis. Photographed at the

Brookfield Zoo, Chicago.
611
www.Ebook777.com
Two views of Cyclanorbis senegalensis, the Senegal soft-shell from north-
central Africa. The plastron clearly shows a relationship to Lissemys.
612
all soft-shells, there are only three claws on each foot. Deraniya-
gala rationalizes this reduction by pointing out that the gait of a
soft-shell is so splayed-out that only the lower three digits make
any contact with the substrate and thus have any need of claws.
The shell openings are completely closeable, the anterior one by
an upward flexion of the front part of the plastron and the hind one
by a pulling up of the femoral flaps together with a slight down
ward flexion of the rear part of the carapace. Body scales are
almost obsolete in Lissemys, but there are five separate crescentic
scales and a circular flat scale on each forelimb, and also a flat
crescentic scale on the sole of each hind foot. The tail is very short
in both sexes.
Lissemys punctata granosa differs from the above in having a
uniformly brown carapace, black longitudinal streaks on the head
(at least in the young), and an entoplastral callosity of moderate
size. (This callosity is called ‘entoplastral’ because it lies in the
same position as the entoplastron of hard-shelled turtles. Soft-
shells lack a true entoplastron). L.p. scutata has the first marginal
bone smaller than the second (the reverse of the condition in punc
tata and granosa), a very large entoplastral callosity, and a brown
carapace spotted or reticulated with black..
In Ceylon, Lissemys is reported to feed on frogs, fishes, Crusta
cea, water snails, and earthworms. It is known to prowl on land at
night, but is usually found in slow moving or still water (rarely in
fast rivers); it often buries itself in soft mud or sand. This turtle is
capable of crushing heavy-shelled molluscs with the flat plates at
the rear of the jaws. The eggs are hard-shelled and spherical,
measuring about 1.2” to 1.3” in diameter. Deraniyagala reports
that several batches of eggs ranging from two to four appear to be
laid within a few weeks during the breeding season. On the other
hand,. Malcolm Smith reports that ten or twelve eggs are laid at a
time. The nests are frequently raided by otters, mongooses, and
jackals.
GENUS CYCLANORBIS
The Senegal soft-shelled turtle, Cyclanorbis senegalensis, is
known from the Sudan west through Cameroon and Gabon to Sene
gal. It has been recorded from Senegal, the Gambia, Chad, Gabon,
Cameroon, Togo, Nigeria, and the Ivory Coast. This species
reaches a maximum length of about fourteen inches. The carapace
of the adult is light brown to blackish-olive, the periphery narrowly
edged with white. The plastron is white, clouded with brown espe
cially on the flaps that conceal the hind limbs. The head and neck
are olive and blackish above, lighter below. The hatchling is grey
ish or brownish, sparsely spotted with black and with indistinct
lighter mottlings, especially along the hind margin of the cara-
613
www.Ebook777.com
An Australian snake-necked turtle from northern Queensland, Chelodina ex-
This is the largest Australian side-neck. Photo by Dr. C.M. Bogert.
pansa.
an Australian snake-necked turtle. Photo by

Chelodina longicollis,
614
At full extension, the neck of Chelodina longicollis is almost as long as the

turtle’s carapace. Photo by G. Marcuse.
615
www.Ebook777.com
pace. The plastron may be quite heavily blotched with dark pig
ment. The head, neck, and limbs are grey or brownish, finely spot
ted with white. There are seven to nine plastral callosities, the
large hyo-hypoplastral pair nearly meeting mesially in large spe
cimens. The callosities become very strong in adults. The cara
pace, which is raised into numerous longitudinal ridges in hatch
lings and has a granulated surface in adults, includes a preneural
bone and a variable number of neural bones, usually at least six.
The neurals do not form a continuous series, but are interrupted in
several places by pleurals meeting mesially. This form of neural
reduction is also displayed by certain specimens of Caretta and
Carettochelys. In Cyclanorbis senegalensis sometimes all of the
first three pairs of pleurals have a median line of contact.
Relatively little is known of the habits of this species. When
captured or handled, it tends to retract its extremities and makes
no attempt to bite. The natural diet has not been recorded, but it
feeds readily on dead fish in captivity. In some parts of Africa
local people keep turtles of this species in their wells, probably to
keep them free of debris and putrefying matter. They are also
caught by the Bari tribesmen of Bahr el Jebel, who use the shells
as basins and the fleshy margins for food. A female specimen
about nine inches long from Duem contained six hard-shelled eggs,
each about 1.4” in diameter. It is possible that this turtle had al
ready deposited part of its clutch, since its cloaca was extremely
flaccid. The hatchling is about 1.8” in length. This is normally a
fluviatile species, but during the Gambian rainy season it is re
ported to move into the marshes.
The Nubian soft-shelled turtle, Cyclanorbis elegans, has a
range that covers a large part of that of the preceding species, and
in some areas the two species appear to be grossly sympatric. C.
elegans is known from the Sudan, Chad, Nigeria, and Togo. It is a
large species, reaching a carapace length of over two feet, and dif
fers from C. senegalensis in many points. Thus there are only two
plastral callosities (a hyo-hypoplastral pair, widely separated) in
most cases (occasionally small xiphiplastral callosities occur
also), the preneural bone is absent, and the neural bones, eight or
nine in number, either form a continuous series or only the last two
are separated. The adult has a light brown to olive-green carapace
with light green spots and vermiculations around the edge. The
plastron is yellow with obscure darker blotches. The hatchling has
a dark olive-brown shell with irregular yellow blotches and a few
small yellow spots around the edge. The head of the adult is dark
brown with light green vermiculations; the temporal region is
olive-green, and the underside is lighter and sometimes speckled.
The head and neck of the hatchling are grey-brown above with
light spots toward the front and passing into a darker shade be-
hind. The forefoot has four sharp-edged crescentic skinfolds on its

upper surface (five or six in senegalensis). The nostril has a papil
la-like projection from the median wall (as do many American
soft-shells), and the tail is extremely short, but without any spe
cific flap to allow for its complete concealment. The carapace of
the hatchling has a moderate vertebral keel and numerous fine
tubercular ridges more or less longitudinally arranged.
The habits of this species are unknown. It is eaten by the Bari
tribesmen.
GENUS CYCLODERMA
The two species of Cycloderma are easily distinguished from
Cyclanorbis species by the more forward location of the eyes; the
postorbital bar is considerably wider than the diameter of the
orbit, which in turn is often wider than the distance between the
orbit and the nasal opening.
Aubry’s soft-shelled turtle, Cycloderma aubryi, is known from
the Congo, Cabinda, and Gabon. The carapace, which may reach a
length of nearly two feet, contains a continuous series of eight or
nine neural bones, only the last pair of pleurals meeting on the
midline. The overall color of the carapace is brown, variable in
shade but with a strong dark brown median line along the anterior
third and with a few scattered spots of brown or black on the rest of
the shell. The plastron in the young is yellow with extensive dark
brown markings. In young specimens there is a vertebral keel and
numerous dorsal tubercles. Plastral callosities are absent in the
hatchling, but seven appear as the turtle grows, and they become
large and almost contiguous in adults. Both femoral and caudal
flaps are present, permitting complete concealment of the rear ex
tremities. The head of Cycloderma aubryi is primarily reddish,
with five longitudinal brown lines, the lowermost of which pass
from the nostrils through the eye to the soft skin of the shoulder
region. The median line is strong and is continued on the carapace.
This median line on both head and shell is unique among soft-shells
but is also found in two West African mud turtles, Pelusios gabon-
ensis and P. niger, at least in the immature stages. No explanation
of this parallelism in coloration of these sympatric species has
been forthcoming. The forelimbs are adorned with six or seven
sharp-edged crescentic skin-folds.
The habits of Aubry’s soft-shelled turtle have not been studied.
Its habitat apparently embraces aquatic situations in rain forests.
The bridled or Zambezi soft-shell, Cycloderma frenatum, is
found in Tanzania, Mocambique, Zambia, and Malawi. Large
adults are nearly two feet long and may weigh as much as thirty
pounds. The shell is longer, narrower, and more convex than that
of aubryi. Usually eight or nine, occasionally seven, neural bones
are present; the last pair, or two pairs, of pleurals meet on the
617
www.Ebook777.com
Plastral view of Chelus fimbriatus. Photographed at the London Zoo.
The matamata turtle, Chelus fimbriatus, a grotesque South American

species. Photo by Dr. Herbert R. Axelrod.
618
Hatchling Geoffroy’s side-necked turtle, Phrynops geoffroanus geoffroanus.

The coloration fades as the turtle matures, but the white line through the eye
is constant. Photo by Dr. Herbert R. Axelrod.
619
www.Ebook777.com
The Zambezi soft-shell, Cycloderma frenatum, of east-central Africa. Notice
the small white spots outlining the dark lines on the neck.
midline. The plastron bears seven very well-developed callosities

in adults. The carapace of hatchlings is unicolor, pale green to
leaden, usually with a white periphery. The surface is somewhat
rough and tuberculate. In adults the carapace is pale to dark olive,
uniform or with a trace of blotching. The plastron of the hatchling
is pale yellow or almost white, usually with a dark blotch on the
umbilicus and a small one in the anal region, and paired blotches
in the humeral and femoral areas of the plastron. The plastron of
the adult is white or pinkish, sometimes (especially in females)
with pearly grey vermiculations. The head of the hatchling is grey
with a dark bar between the eyes and five longitudinal lines, some
times wavy and broken, from the occipital region to the base of the
neck. In adults these lines may almost disappear and the head and
neck may display numerous white dots.
620
The eggs are laid from December to March. Recorded

clutches numbered between 15 and 22 eggs, hard-shelled and 1.2 to
1.4 inches in diameter. The Zambezi soft-shell lives in rivers,
lakes, and stagnant ponds. The turtles become active during rainy
weather, and it is at such times also that hatchlings emerge. The
species appears to be shy and inoffensive, though equipped with
the usual strong jaws and active disposition of soft-shells.
This is a carnivorous species, feeding mainly on meat or fish in
captivity, but apparently subsisting mostly on molluscs in the wild
state. In Lake Malawi the turtles use their powerful forelimbs to
dig for buried mussels.
GENUS TRIONYX
The New World soft-shells are grouped into only four full spe
cies, distributed from southern Canada to northern Mexico. They
are all referable to the genus Trionyx and are characterized by
being of medium size (adult females around 12 to 15 inches long),
having no true carapacial ocelli at any stage of development
(though small circular spots may be present), and in showing a
strong tendency toward reduction or complete elimination of the
eighth pair of pleural bones.
Trionyx ferox, the Florida soft-shell, is the largest species.
The greatest length is usually stated as eighteen inches, but I have
personally measured a 20” specimen, and it is possible that still
greater lengths may sometimes be reached. This species is found
almost throughout Florida, though it is absent in the Keys and in
the western part of the Panhandle. It also occurs in the coastal
plain of southeastern Georgia and in southern South Carolina.
The coloration of the young of Trionyx ferox is very distinc
tive. The hatchling is about 1.5” long and has a yellowish-olive car
apace marked with very large, slaty-grey spots. Some of these
may be light centered, and around the edges of the carapace the
spots may be so closely packed that they become confluent. The
rim of the carapace is marked with a narrow yellowish line. This
light border is visible from below, but otherwise the ventral sur
faces of the shell are slate-grey with only indistinct light mottling
on the anterior parts. The upper surfaces of the extremities are
olive with yellow and orange markings which form indistinct
stripes on the head, upper jaw, and neck. At this stage the cara
pace is adorned with fine longitudinal tubercular wrinkles which
grade into coarse rounded knobs in the nuchal region. There is a
light tubercular skin fold extending from the heel to the base of the
tail on each side, a single sickle-shaped scale on each heel, and
three such scales on the outer face of the forelimbs.
The striking juvenile coloration of young Florida soft-shells is
lost with age, the first stage being a lightening of the dark plastral
621
www.Ebook777.com
Lateral view of Phrynops geoffroanus geoffroanus swimming. Photo by
South American gibba turtle, Phrynops gibba, from the Upper Commewijne
River, Surinam.
622
South American toad-headed turtle from Colombia, Phrynops nasuta

wermuthi.
South American toad-headed turtle, Phrynops nasuta, a side-necked turtle

with an unusually large head. Photographed at the Bronx Zoo.
623
www.Ebook777.com
pigmentation. Adults have somewhat elongate brown carapaces
on which traces of the juvenile dark markings may or may not be
retained. The longitudinal wrinkles and anterior rounded tuber
cles, however, are retained. The age, or at least size, at which the
juvenile coloration is lost is very variable, some four-inch speci
mens having the typical adult coloration and some eight-inch spe
cimens retaining the juvenile pattern. The head stripes are lost in
the adult, the head and neck becoming uniform brown, grey, or
pinkish. The head is relatively broad in this species, compared to
that of the other American soft-shells, and the jaw surfaces often
become greatly enlarged, especially in old males. Females reach
a considerably larger size than males.
The Florida soft-shell is less fluviatile than its relatives, and in
those areas where it is grossly sympatric with the spiny soft-shell
(Trionyx spinifer), it tends to be found in still water while the other
species frequents rivers and streams. It is common in the Okefe-
nokee Swamp and is also very plentiful in the Florida Everglades
which, while technically a river, is from a turtle’s point of view
Juveniles of the Florida soft-shell, T r i o n y x f e r o x , showing the distinctive

large brownish spots on the carapace. Photo by R.J. Church.
624
more like a swamp. The species is sometimes found in the sluggish

lower reaches of rivers and has been recorded in brackish water.
This is primarily a carnivorous species, eating large quanti
ties of invertebrates. Mature males have jaw modifications that
suggest an inclination toward eating molluscs. Small quantities of
fish are eaten, but it is probable that most if not all fish eaten were
already dead or incapacitated.
Eggs are laid from March to July in Florida, but apparently
not earlier than June further north. Females emerging from the
water to nest are cautious, holding the head high and returning to
the water if disturbed. The nest cavity is made with the hind feet
and usually in sandy soil or pure sand; many nests in Florida are
made in ruts or on the edges of little-used dirt roads. Recorded
clutch sizes range from 7 to 22 spherical, hard-shelled eggs each
about one inch in diameter.
The Florida soft-shell is a popular food item in many parts of
its range, to the extent that some populations in northern Florida
have declined markedly in the last few years.
The Florida soft-shell shows a number of characteristics, not
shown by the other American soft-shells, that suggest close rela
tionship with some of the Old World species. Among these features
are the relatively frequent occurrence of an eighth pair of pleural
bones (in about one individual out of three), the comparatively
large size, tolerance of brackish water conditions, and the longitu
dinally wrinkled carapace. Since the soft-shells are thought to
have originated in the Old World, it is likely that T. ferox, with its
various Old World characteristics, is the most primitive New
World species. It appears likely that ferox, or the populations
directly ancestral to modern ferox, was once much more widely
distributed to the north and west and that it was forced to retreat to
Florida and the lower coastal plain by increasingly cold conditions
in the Tertiary. It will be recalled that the same phenomenon is
postulated to have caused the retreat of the Florida snapping tur
tle, Chelydra serpentina osceola, to Florida, and of the closely re
lated Mexican snapper, C. s. rossignoni, to Veracruz, thus splitting
the range of the ancestral population into eastern and western Gulf
enclaves. It is possible that the retreat of the ancestor of Trionyx
ferox also left a population on the western side of the Gulf which is
now restricted to a small area of central Coahuila. This population
today is considered to constitute a full species, Trionyx ater, the
Mexican black soft-shell.
Trionyx ater is only found in the basin of Cuatro Cienegas,
Coahuila, Mexico, where it has been caught in Tio Candido Pond,
16 km. south of the village of Cuatro Cienegas, and Anteojo Pond, 9
km. west of the Cuatro Cienegas. Both ponds are permanent and
have clear water. It is interesting to note that the box turtle Terra-
625
www.Ebook777.com
Twist-necked turtle, Platemys platycephala, from Guyana. Photo by
Argentine snake-necked turtle, Hydromedusa tectifera. Photographed at the

American Museum of Natural History.
626
Baby Carettochelys insculpta from New Guinea, showing the serrations on

the carapace typical of the young of this species.
627
www.Ebook777.com
pene coahuila is similarly confined to the Cuatro Cienegas Basin.
Trionyx ater shows ferox-like characteristics in having a broad
head, short snout, overall dark dorsal coloration in adults, similar
dorsal limb coloration, reduction and blunting of the anterior cara
pace tubercles, and a still-water rather than fluviatile habitat. It
is, however, a small species not known to exceed a length of about
ten inches and shows some points of similarity to both Trionyx
spinifer emoryi and to T. muticus. It is probable that T. ater,
though today reduced to an insignificant population, is a relatively
unchanged descendent of the ancestral stock which gave rise to all
American soft-shells.
Apart from the characters mentioned above, T. ater is charac
terized by frequent fine corrugation of the posterior carapace mar
gin, especially in females, and reduction of the septal ridges in
adult males. Males have a pale outer carapace rim, a ‘sand-
papery’ texture of the hind part of the carapace, a broadened
snout, dark grey or slate dorsal coloration with small whitish dots
toward the rear, and no contrasting pattern on the soft parts. Fe
males lack the pale carapace rim, having an overall contrasting
dorsal pattern of blackish and slate-grey, and have many blackish
flecks and markings on the underside of the carapace and plas
tron.
The most widespread soft-shell species in the New World is the
spiny soft-shell, Trionyx spinifer, whose range extends from south
ern Ontario and Quebec through northwestern Vermont and wes
tern New York south to south Georgia and to the Florida pan
handle, and east to central Montana, Wyoming, and Colorado, and
southern New Mexico. In Mexico it occurs in northern Coahuila,
Nuevo Leon, and Tamaulipas, and northeastern Chihuahua. It is
found in many parts of the Colorado River drainage in Arizona and
the neighboring parts of California, Nevada, and New Mexico, but
these populations are thought to be introduced.
The spiny soft-shell receives its name from the conical projec
tions along the anterior margin of the carapace, though these are
more defined in some races than in others. The carapace has a uni
form sandy or brown ground color without the large markings
typical of T. ferox, but with many small dark spots or ocelli in the
eastern races and small white spots in the western forms. These
are fluviatile turtles for the most part, very wary and fast, of pred
atory and scavenging habits, and fond of burying themselves com
pletely in sandy river bottoms. Six subspecies are currently recog
nized. T. s. spinifer, the eastern spiny soft-shell, is found east of the
Mississippi River and south as far as extreme northern Alabama
and west-central Mississippi. It is absent in the Atlantic drainage
systems except probably for the Hudson-Mohawk drainage. T. s.
hartwegi, the western spiny soft-shell, occupies the range of the
628
A well-marked eastern spiny soft-shell, Trionyx spinifer spinifer. The

subspecies of this species are often hard to distinguish. Specimen from the
Lantz collection.
species west of the Mississippi and north of central Louisiana and

central Oklahoma. T.s. asper, the Gulf Coast soft-shell, is found
from southern North Carolina through almost all of South Caro
lina, Georgia, Alabama, and eastern and southern Mississippi.
T.s. pallidus, the pallid spiny soft-shell, is found in southern Okla
homa, eastern Texas, extreme southwestern Arkansas, and the
western half of Louisiana. T.s. guadalupensis, the Guadalupe soft-
shell, is found in south-central Texas in the Nueces and Guadalupe-
San Antonio Rivers. T.s. emoryi, the Texas spiny soft-shell, is
found in the Rio Grande drainage in Texas, New Mexico, and
northern Mexico; in the Rio San Fernando and Rio Purificacion
drainages in northeastern Mexico; and the Colorado River drain
age in Arizona, New Mexico, and southern Nevada.
The eastern spiny soft-shell, T.s. spinifer, is recognized by the
carapace pattern which in juveniles and adult males is made up of
numerous relatively large and well-defined black ocelli; also only
629
www.Ebook777.com
Fly River turtle, Carettochelys insculpta, from southern New Guinea. This un
common species is highly aquatic and has limbs that are modified into flip
pers. Photographed at the Bronx Zoo.
630
Female Podocnemis expansa excavating its nest. Photo by Harald Schultz.
Copulating loggerhead turtles, Caretta careta. The loggerhead is the most

common sea turtle in the United States. Photographed at Marineland of
Florida.
631
www.Ebook777.com
a single dark marginal line separates the pale rim of the carapace
from the dorsal ground color. The ocelli nearest the middle of the
shell are the largest. Large females lose the clear juvenile pattern,
the ocelli becoming broken and obscure and the background color
ation blotched rather than uniform. The snout is adorned with pale
dark-bordered stripes that join at an acute angle in front of the
eyes. Pale yellow black-bordered stripes are continued backward
from the eye and also from the corner of the mouth. The upper face
of the front limbs is adorned with a complex pattern of dark and
light curved stripes. Adult males have small conical tubercles on
the anterior part of the carapace; in large females blunt tubercles
are present on the middle of the carapace as well as the front.
Juveniles of T.s. hartwegi are distinguished from those of spin-
ifer by the smaller dorsal ocelli or by the presence of small black
spots instead of true ocelli. The east to west decrease in ocellus
diameter is somewhat clinal, however, and the Mississippi River
is rather arbitrarily designated as the dividing line between the
races. In hartwegi only a single marginal line is present. The head
is narrower than in asper, guadalupensis, and emoryi, and the
carapace is wider than that of emoryi and reaches its widest point
about half way back.
The Gulf Coast soft-shell, T.s. asper, has black ocelli and spots
over the carapace, and two or more distinct marginal lines. These
markings, however, are only really distinct in juveniles and
males. The striping on the snout is variable and sometimes com
pletely absent, but when present the head stripes do not meet on
the neck as they do in the other subspecies.
632
T.s. pallidus has a pattern of white spots in certain individ

uals ; these are usually more clearly defined in young specimens
and tend to be concentrated on the anterior third of the carapace.
The pale carapace rim is less than four times wider posteriorly
than laterally.
T.s. guadalupensis usually has white dots all over the cara
pace ; in adult males the white dots may reach 3 mm in diameter
and be encircled with blackish ocelli.
The subspecies emoryi, as currently defined (before 1962 tur
tles now called pallidus and guadalupensis were included in this
race), has a juvenile pattern of white dots which are not encircled
by dark ocelli and which are confined to the posterior third of the
A specimen of Trionyx spinifer emoryi from the Rio Verde, San Luis Potosi.
carapace; the pale rim of the carapace is conspicuously widened,

being four to five times wider posteriorly than laterally; there is a
dark triangle in front of the eyes, the base line of which connects
the anterior margins of the orbits; and the pale postocular stripe is
interrupted, leaving conspicuous pale, usually dark-bordered
blotches behind each eye. This is the most variable of the sub
species. Details of the color pattern in different areas and of vari
ant individuals are given by Webb (1962). Several thousand free-
633
www.Ebook777.com
The most popular aquatic pet turtles in the world were juveniles of
Pseudemys, Chrysemys, and Graptemys species from the southeastern
United States. Young individuals intended to be sold as pets were harvested
by the millions annually and shipped all over the world. This is a young
Pseudemys concinna hieroglyphica. Photo by Dr. Herbert R. Axelrod.
Tortoises like the European Testudo shown here are comparatively more
popular as pets in Britain and Germany than in the United States. Photo by G.
Marcuse.
www.Ebook777.com
living soft-shells of this race are to be found in the moats and
waterways which separate the exhibits in the Gladys Porter Zoo in
Brownsville, Texas.
In general, males of the spiny soft-shell are smaller than fe
males; Conant found that of 71 specimens of spinifer from Ohio,
the largest female was 14.4 inches long and the largest male only
6.50 inches. On the other hand, a male of emoryi from Houston,
Texas, measured 13.9 inches in length. The greatest length at
tained by this species is about eighteen inches. Webb estimated
that turtles of this size are about sixty years old.
The average clutch size for northern populations of spinifer is
about 20 eggs; only one clutch is laid in a season. The eggs are
spherical, with thin, brittle shells, and average 1” to 1.2” in
diameter. Oviducal shelled eggs are slightly smaller on the
average than eggs taken from nests, and it appears likely that
some post-deposition swelling takes place. The normal incubation
time is sixty to seventy-five days, though the young from late-
season nests may winter underground. Nesting usually takes place
in June and July.
The spiny soft-shell is a predominantly carnivorous species;
the food consists mainly of insects and crawfish, though a small
quantity of fish is eaten and some individuals evidently take to a
predominantly shellfish diet, judging by the massive flat jaw
plates that sometimes develop. It is possible that occasionally a
very large individual will eat a duck or duckling, pulling it under
by the feet from below. In some areas appreciable quantities of
vegetation are eaten, and it is said that soft-shells may be trapped
in numbers using traps baited with watermelon rind. Spiny soft-
shells are considered excellent eating by cognoscenti in most
places where they occur, but there are few places in the United
States today where they could be considered an important item of
commerce.
The smooth or spineless soft-shell, Trionyx muticus, is wide
spread in the central United States, being known from extreme
western Pennsylvania west to southeastern South Dakota, south to
northeastern New Mexico and the Gulf Coast from about the mid
point of the Texas coast east to the tip of the Florida Panhandle.
This species, like certain of the races of spinifer, shows some evi
dence of being derived from an afer-like ancestor. The size is rela
tively small (the record length is 14 inches, but the usual adult size
is much less than this), and the carapace is smooth and devoid of
tubercles even in the nuchal area. The septal ridges, which are
well-defined in spinifer but reduced in male ater, are absent in
muticus. The broad juvenile carapace is retained in the adult, and
the widest point is about half way along the carapace (as in the
m lire eastern and northern races of spinifer). The skull is distinc
tively elongate and narrow, with large orbits and a long snout.
636
Two subspecies of Trionyx muticus are recognized: muticus

from the Mississippi drainage from extreme western Pennsyl
vania, southern Minnesota, and South Dakota south to Tennessee,
Louisiana and Oklahoma; also in streams of the Gulf coast drain
age from the Mississippi in Louisiana westward into Texas includ
ing the Colorado River drainage. Calvatus ranges from eastern
Louisiana north of Lake Pontchartrain east as far as the Escambia
River in the extreme western tip of the Florida Panhandle. The
race occurs as far north as northeastern Mississippi and probably
also northern Alabama, though precise localities in Alabama are
still lacking. In the typical race there is a juvenile pattern of dusky
spots and usually short lines or streaks. There is a very narrow
light band around the edge of the carapace. Ill-defined pale stripes
are usually evident on the snout just in front of the eyes, and also
there is a pale postocular stripe with narrow or indistinct dark
borders. In calvatus there is a juvenile pattern of large circular
spots, often ocellate; there are no stripes on the dorsal surface of
the snout; the pattern on the dorsal surfaces of the limbs is of fine
markings which do not contrast with the ground color; and the
pale postocular stripes have thick black borders approximately
half the width of the pale stripe in adult males.
Both races may be distinguished from the subspecies of spini
fer not only by the characters mentioned above, but also by the
obtusely-angled epiplastron (approximately right-angled in spini
fer), the relatively large plastral callosities in adults, and the
wider hyo-hypoplastral bridge. The hatchlings are tiny, flat, insub
stantial little creatures which average about 1.35” in carapace
length and often have a distinct pink tinge to the shell.
Trionyx muticus is a highly fluviatile species, being found in
rivers and streams but almost never in still water. This may re
flect less of a tendency toward overland wandering than is shown
by spinifer, and this tendency is also manifested in the choice of
nesting sites; spinifer may frequently wander some distance over
land before excavating its nest, while muticus usually nests on
mid-river sandbanks.
Males of muticus reach maturity at shell lengths of only 4.5 to
5 inches, but females reach maturity at 7 to 9 inches. Nesting takes
place from late May to late July. The nest is six to nine inches
deep, and the eggs are arranged in this cavity with the hind feet as
they are laid. Recorded clutches number between 4 and 33 eggs,
but around twenty is usual. As with certain sea turtles, larger tur
tles lay more eggs than smaller ones. The eggs are spherical and
0.8” to 0.9” in diameter. The incubation period is said to be 65-77
days.
Ernst and Barbour report that the food of muticus consists of
fish, frogs, tadpoles, mudpuppies, crayfish, aquatic insects, snails,
637
www.Ebook777.com
The American box turtles such as this male Terrapene Carolina Carolina are
hardy, interesting, and completely inoffensive, and they make good pets, but
their ownership by private individuals is in some localities prohibited by law.
Photo by Dr. Herbert R. Axelrod.
Turtles have played a part in the cultures of many people of the world, and
they have obtained good representation in artifacts. This figurine was pro
duced by an Amazonian Indian. Photo by Harald Schultz.
638
Many conservationists and animal welfarists have condemned the turtle

trade as dangerous both to natural turtle populations and, because of
disease, to turtle owners. Today the trade in small aquatic species is effec
tively stopped, but fanciers still manage to successfully keep various
species.
639
www.Ebook777.com
bivalves, and worms. In addition, algae, fruits, and hard seeds and
nuts may be eaten.
The single Trionyx species in Africa is the large and wide
spread Trionyx triunguis, both of whose names refer to the univer
sal soft-shell feature of three claws on each foot. The Nile soft-
shell, as this species is often called, is found in Egypt, the Sudan,
Ethiopia, Somalia, Uganda, Kenya, Angola (including Cabinda),
Zaire, Gabon, Cameroon, Togo, Ghana, Liberia, Sierra Leone, the
Gambia, Senegal, and Portuguese Guinea—in fact the whole of
Africa except for the Northwest and the South. It occurs along the
entire length of the Blue Nile, but in the White Nile it is not found
above the Murchison Falls in Uganda. Thus it is found in Lake
Rudolph and in Lake Albert, but not in Lake Victoria.
The Nile soft-shell is a very large species, the largest individ
uals being some three feet in carapace length and a hundred
pounds in weight. The skull is somewhat flatter and more elongate
than that of the American soft-shells. The overall color of the ani
mal is dark brown or olive profusely speckled on all dorsal sur
faces with small yellow spots; these spots may become obscure or
even disappear in old adults. The underside may be immaculate
white or there may be dusky vermiculations on the anterior of the
plastron, and there may also be light, dark-bordered spots on the
underside of the dorsal disc. The habitat of this species typically
embraces ponds, lakes, and rivers. There is an old record for a
large specimen having been caught at sea two or three miles from
the mouth of the Gaboon River, and it is reported to be found off
the coast of Senegal near Dakar quite frequently. It has been
known for a long time that soft-shells may also be swept out of the
mouth of the Nile and be deposited on the shores of Syria and
Israel. However, I was surprised to be informed recently by Dr.
Ralph Hathaway that the species is found on the coast of Turkey,
not as rare strays incapacitated by immersion in sea water, but as
a population of strong, vigorous animals living in 6 to 15 fathoms of
sea water with a salinity of at least 38 parts per thousand. More
over, the animals are plentiful enough to be a pest along the entire
coast; fishermen find them so vicious they do not like to bring
them aboard. Where these marine soft-shells breed is unknown;
possibly they nest in the same sandy beaches that are used by
green turtles and loggerheads. I once picked up a dried carapace
of a soft-shell about six inches long on the shore of the Gulf of
Iskenderun in southern Turkey.
In Egypt, the Nile soft-shell is reported to nest in April, the
eggs numbering 50 to 60 and being buried in sandbanks. Reports
for other areas suggest that sometimes as few as 25 eggs are laid.
The young have indistinct median keels and carapacial tubercles
arranged in wavy longitudinal lines. Plastral callosities are absent
640
in the young, but four develop as the animal grows—a hyo-

hypoplastral pair and a xiphiplastral pair.
Trionyx triunguis appears to be an omnivorous species, repor
ted stomach contents ranging from molluscs, frogs, fish and in
sects to palm nuts and dates. The species is used for human con
sumption in many parts of its range.
The Euphrates soft-shell, Trionyx euphraticus, is smaller than
the Nilotic species, being adult at about fifteen inches. The dorsal
coloration is dark green, often with small light spots or irregular
light and dark blotches. The undersurfaces are white. The eighth
pair of pleural bones is somewhat reduced, being not more than
half the size of the seventh pleurals. The proboscis is relatively
short, and the distance between the snout and the orbit is about
equal to the diameter of the orbit and about three times the width
of the postorbital bar. There is no median ridge on the mandibular
symphysis. There is a pair of feebly-developed plastral callosities
located on the hyo-hypoplastra.
This species appears to be abundant in at least parts of Syria
and Iraq and is probably little molested by local people. Never
theless, it is very rarely seen in zoos or in private collections, and
its habits have not been described.
The Euphrates soft-shell, Trionyx euphraticus, of Syria and Iraq. Photo by

Dick Bartlett.
641
www.Ebook777.com
For several hundred miles to the east of the range of euphrati-
cus, no soft-shells of any species are found. However, the large
Ganges soft-shell, T. gangeticus, is also found in the Indus River of
Pakistan, where it ranges north at least as far as Dera Ismail
Khan and Montgomery. It is also found in the Mahanadi River sys
tem in Orissa, and in the Ganges it is known from extreme eastern
India, Bangladesh, and southern Nepal. Adults of both sexes com
monly reach a length of eighteen inches, and the maximum shell
length is probably around 27 inches. The shell, as in most soft-
shells, is low and broadly oval in outline; callosities are not partic
ularly evident in living specimens, but are present on the bones of
Detail of the head of the Ganges soft-shell, Trionyx gangeticus.
the carapace and on four areas of the plastron. The head is rela
tively large and broad, with dorso-laterally situated eyes and a
relatively thick proboscis. The jaws have expanded, granular
crushing surfaces. The tail is short in both sexes. Adults are dull
olive or greenish dorsally with vague dark reticulations; the plas
tron is ivory-white, slightly darker on the callosities. Juveniles are
greenish or buff-colored dorsally with a complex reticulate pattern
enclosing four rather irregular ocelli on the carapace and strong
oblique black lines reaching postero-ventrally from the eye and
the top of the head. There is also a longitudinal black line along the
top of the head. The iris of the eye is light greenish-yellow.
642
The Ganges soft-shell is a highly aquatic species living in deep,

turbid rivers, but it is frequently seen basking on sandbanks or sit
ting in shallow water with its head sticking out of the water. In diet
it appears to be largely herbivorous, eating aquatic vegetation,
though animal matter has been found in some stomachs. In Pakis
tan nesting takes place in October, when the waters are receding.
The eggs are spherical, hard-shelled, and about 114” in diameter.
The flesh of this turtle is highly popular for food, and large num
bers are sold in Calcutta for this purpose. A large collection of
Ganges soft-shells is kept in an enclosed pool at Puri, in Orissa, on
part of the Mahanadi River system, where the Mohammedans
maintain them for religious purposes. The pool has an area of
about half an acre, and the turtles have apparently lived there for
centuries, climbing out on the banks regularly to lay their eggs.
Annandale considered these turtles to represent a distinct sub
species (mahanaddicus), but Malcolm Smith dismissed this pos
sibility. It is in any case a questionable procedure to base a new
subspecies on a highly inbred and long-term captive population.
Another widespread Indian species is the Nagpur soft-shell,
Trionyx leithi, known from the Ganges and rivers of peninsular
India as far south as Madras. This species appears to be quite sim
ilar to gangeticus, but the bony snout is longer than the diameter of
the orbit (about the same length in gangeticus), and the inner edge
«he mandible is not raised, the mandibular symphysis being flat.
o, the carapace of the young is adorned with four rather well-
.lined ocelli, or a pattern of eye-like concentric circles; one or
two additional ocelli of smaller size may also be present. The head
markings are generally similar to those of gangeticus, though the
black lines may be broken up. The maximum recorded shell length
is around twenty inches.
The third Indian species is the so-called peacock soft-shell,
Trionyx hurum, which is found in the lower reaches of the Ganges
and Brahmaputra. Adults, which may reach a carapace length of
about two feet, are not strikingly marked, but the young are beau
tiful animals, with dark olive-green carapaces reticulated with
black and with four (or occasionally more) very striking eye-spots
or ocelli. Each ocellus is composed of a black central disc narrow
ly edged with a light yellow band, this in turn being circled by a
broad dark band (though not as intensely dark as the center),
while the outermost circle is the broadest and is olive-green in
color. The carapace is also adorned with a narrow yellow rim and
numerous broken longitudinal ridges. The head is dark green or
black with numerous yellow spots, mostly small but a few of them
very large; one of the enlarged spots is across the base of the
snout, one behind each eye, and others behind each comer of the
mouth. As the turtle grows, the carapace ocelli and yellow head
643
www.Ebook777.com
spots become less distinct, and the dorsal surfaces become gener
ally darker and the undersurfaces paler.
The snout of T. hurum is more strongly downturned than that
of its close relatives; the snout is longer than the diameter of the
orbit, which in turn is two or three times the width of the postorbi
tal bar. The mandibular symphysis has a median groove that is
considerably longer than the orbital diameter.
The function of the dorsal eye-spots of this and several other
Oriental soft-shells is still uncertain, but it is most likely that they
serve the purpose of giving potential predators a momentary scare
when they suddenly come into view as the turtle seeks refuge; the
predator’s hesitation before it realizes the ruse may well be suffi
cient to allow the turtle to escape. It might be argued that four
staring eyes does not mimic any possible animal that might
frighten the predator, but this is probably a case of what is known
as a ‘supernormal stimulus.’ If the sudden appearance of two big
staring eyes is scary, then the sudden appearance of four such
eyes is even more scary. As the turtle grows and the number of
potential predators decreases, it is permissible for the dorsal pig
mentation to become disorganized, thus causing the ocelli to dis
appear.
The next species, Trionyx nigricans, apparently no longer
exists in the wild state, but a semi-captive colony has persisted in
an enclosed pond, or tank, attached to the Mahommedan shrine of
Sultan Bagu Bastan about five miles from Chittagong, Bangla
desh. The turtles were reported by Annandale in 1912 and 1914 to be
so tame that they would come to feed when called. Otherwise they
spent the daytime half buried in the mud at the bottom of the pond,
while nights were spent out of the water on a small hill. Oviposition
took place in rainy weather, the eggs being laid in this same hill.
Despite the recent natural disasters and social upheavals in Bang
ladesh, this colony is still extant; it was recently visited by Guy
Mountfort of the World Wildlife Fund, who took photographs of the
turtles being fed by hand.
Trionyx nigricans is a large species, the largest individuals
being about 32 inches long. The structure of the carapace and plas
tron is essentially similar to that of T. gangeticus, but the alveolar
surface of the mandible is not raised at its inner margin and the
length of the median mandibular ridge equals the orbital diameter
(less than the orbital diameter in gangeticus). The coloration of
the adult was described by Anderson, who described the species in
1875, as follows:
“Carapace dark blackish plumbeous, with a tinge of

olive due to the presence of blackish spots, among which are
intermixed many rusty brown spots, which overlie as it were
644
The captive
population of
Trionyx nigricans
at Chittagong,
Bangladesh. This
is the only known
surviving popula
tion. Photo by
Guy Montfort
the black spots. The head, neck, and upper surface of the
limbs are almost black; the upper lip in its two posterior
thirds is white; and there is a great white blotch over the
ear. The area between the neck and the forelegs is whitish;
and there are some white spots on the margin of the cara
pace. The head is reticulately spotted; and there is a distinct
infrapraeorbital band, and a trace of another above the
eyes; but the head is so black that these markings are diffi
cult to distinguish. The undersurface of the head and neck is
almost black; and the plastron is densely spotted with black
ish purple, especially over the bones, and the tail is similarly
marked. The claws are yellow. ”
Annandale, who visited the shrine almost forty years after
Anderson, described the turtles as follows:
www.Ebook777.com
“Carapace copper brown indistinctly marbled with a
darker shade; head and limbs apparently dark clay color,
but always covered with mud; top and sides of head bright
glaucous green, taking a yellowish tinge above the eyes and
nostrils and boldly reticulated with black or dark green; in
all very old turtles the coloration of the head might be de
scribed as black with greenish spots which tend to disappear
with age. ”
One small specimen noticed by Annandale had traces of dorsal

ocelli.
The Burmese soft-shell, Trionyx formosus, is known from the
Irrawaddy, Sittang, and Salween Rivers in Burma, and the range
extends close to the frontier with Yunnan Province, China. This
species differs from those discussed so far in that it lacks the ‘pre-
neural’ bone, a laterally expanded bone situated between the
nuchal and the first neural, between the anterior halves of the first
pleural bones, in most soft-shells. The carapace, which in adults
reaches a length of about fifteen inches, is adorned in the young
with four well-defined ocelli essentially similar to those of T.
hurum. The head and neck of the young are ornamented with yel
lowish dark-edged spots, an elongate one on either side at the back
of the head and smaller ones on the temple, at the angle of the
mouth, and on the chin. The head of the adult is yellow with black
reticulations or spots above and on the sides. In the adult the juve
nile ocelli are reduced to obscure black areas. The length of the
snout is not greater than the diameter of the orbit, which is two or
three times the width of the postorbital arch. There is a longitudi
nal median ridge on the mandibular symphysis.
The Burmese soft-shell is reported to be not uncommon in the
lower reaches of the Irrawaddy, but like all Burmese turtles, its
habits are virtually unknown and live specimens are not known in
western collections. However, these turtles are considered sacred
by certain Mahommedans, who maintain a number of them alive
in a tank in Mandalay. Many turtles in this tank are deformed, pos
sibly through excessive inbreeding. Some are much larger than
any known preserved specimens. The custodians prohibit the
removal of the turtles for scientific purposes. They are very tame
and come when called; they are reported to feed on chicken and
rice.
Related to the Burmese soft-shell in that it too lacks the pre-
neural bone is the black-rayed soft-shell, Trionyx cartilagineus.
This is a large species, up to about 28 inches in shell length, found
in southern Burma, Thailand, southern Laos, Cambodia, North
and South Viet Nam as far north as the Gulf of Tonkin, the Malay
Peninsula, Sumatra, Java, and Borneo.
646
The black-rayed soft-shell has a bony snout longer than the

diameter of the orbit and a median ridge on the mandibular sym
physis. The dorsal coloration is very variable, but typical young
specimens have a basically olive-green carapace decorated with
broad black radiating streaks, the entire surface being heavily
speckled with yellow and with one or two irregular black spots out
lined in yellow. However, in many juveniles the pattern is not as
clearly defined as this description would suggest, and in adults the
carapace becomes more or less uniform in color.
The plastron is greyish-white in the juveniles, and the head is
dark brown or grey with numerous small yellow spots. As the tur
tle grows, the head pattern becomes a black reticulation on an
olive ground. The plastron is white in adult males, grey in females.
In some areas the top of the head is adorned with black streaks
that may form an X-shape.
The black-rayed soft-shell is reported to inhabit muddy slow-
flowing rivers, but it is also found in hill streams. Its eggs are de
posited in holes in mud banks. The eggs are 1.2” to 1.4” in diame
ter, but the normal clutch size is not known. The flesh of this spe
cies is considered inferior in flavor to that of most soft-shells. It is
reported to feed upon fish, crustaceans, and amphibians.
The Malayan soft-shell, Trionyx subplanus, is found in Burma,
Thailand, the Malay Peninsula, Sumatra, Java, and Borneo, as
well as on many offshore islands, such as the Mergui Archipelago,
off Tenasserim, or Singkep, in the Riau Archipelago. This is a fair
ly distinctive species which is often placed in a separate genus
(Dogania). It has a large head, an exceedingly narrow postorbital
bar, no preneural bone, and no median ridge on the mandibular
symphysis. The eight pleural bones are all separated along the
midline by the neurals; in most soft-shells the last pair have a
median line of contact. This is one of the smallest of the soft-shells,
fully grown adults being about ten inches long.
The carapace in the young of T. subplanus is marked with
numerous longitudinal wrinkles, especially toward the rear, and is
of a rich chestnut-brown color with fine black vermiculations.
There is a strong black median stripe and two (or three) pairs of
rather small black ocellar markings. The chestnut-brown color
and the black median stripe are continued on the head and neck,
while the sides of the neck are pink to bright red. There are two
oblique black streaks on each side of the black line on the head,
and the chin has black vermiculations. The plastron is pale yellow.
The Malayan soft-shell tends to avoid the large, slow rivers
inhabited by its bigger relatives, preferring hill streams, where it
frequently hides under rocks and flat stones.
Soft-shelled turtles are abundant in many parts of China, and
numerous species have been described. The most prolific author in
647
www.Ebook777.com
this field, a French missionary by the name of Father Heude, de
scribed no fewer than thirteen species of Chinese soft-shells in
1880, which he referred to no fewer than nine genera ( Yuen, Psilo-
gnathus, Temnognathus, Gomphopelta, Coelognathus, Tortister-
num, Ceramopelta, Coptopelta, and Cincistemum). Nevertheless,
all these names are now considered invalid and, apart from Pelo-
chelys bibroni (described below), all Chinese soft-shells are now
referred to only two species, Trionyx sinensis (with two subspe
cies) and T. steindachneri.
The Chinese soft-shell, or suppon, Trionyx sinensis, is known
from the eastern seaboard of China, from Changnam Province in
the south to Hopei and Jehol in the north. It is also found on both
Hainan and Taiwan, and inland has been recorded from the Prov
inces of Suiyuan, Shensi, Kansu, Szechwan, Yunnan, Kweichow,
and Kwangsi. It is known from Korea, Manchuria, North Viet
Nam, and Japan. It has been introduced to some of the Hawaiian
Islands, though only the island of Kauai appears to have an estab
lished population. Records for Singapore, Thailand, the Batan
Islands, and California are considered to be based on introduced
specimens, and it is very likely that a soft-shell turtle from Timor
described under the name Trionyx cartilagineus var. newtoni in
1897 is based on an introduced population of sinensis.
As one might expect for a species which has been given so
many different names, Trionyx sinensis is a very variable species.
Clifford Pope describes the coloration as:
“Extremely variable but generally olive above with or
without dark vermiculations, light yellowish below; large,
dark ventral and small, black dorsal spots usually evident in
juveniles, sometimes persistent to early maturity; head
with or without fine black lines radiating from eyes; throat
light or vermiculated, or dark with yellowish spots. ”
Dorsal ocelli are never present.
Two subspecies of Trionyx sinensis are recognized: T.s. sinen
sis from the more southern part of the range, and T.s. tuberculatus
from Yunnan, central China, and Taiwan. The latter is distin
guished by the discrete, though linearly aligned, tubercles on the
carapace; the typical form has irregular longitudinal ridges on the
carapace rather than tubercles. Also, the plastral spots of tuber
culatus are smaller and less distinct, and there is an area of coarse
tubercles at the base of the neck.
Trionyx sinensis is found in rivers, lakes, canals, and creeks in
China, but does not ascend into cascading streams, although it
may reach considerable altitudes in northern and western China.
It is known to feed upon fish, crustaceans, grasshoppers, molluscs,
and some plant material. Eggs are known to be laid in June in
648
Trionyx sinensis tuberculatus. This specimen of the Chinese soft-shell is pro

bably from Taiwan.
northeastern Kansu. The eggs are spherical and 0.8” to 1” in diam

eter; two to four clutches, each numbering 17 to 28 eggs, are re
ported to be laid annually. Captive females in a Japanese turtle
farm reached maturity when about six years old; copulation takes
place in spring at the surface of the water. Hatchlings are margin
ally over an inch in length. This species is heavily used for food in
most parts of its range; the small introduced population in Hawaii
is exploited intensively.
Trionyx swinhoei (or Trionyx swinhonis), from Shanghai, is
customarily synonymized with T. sinensis, but both Boulenger and
Loveridge and Williams consider it recognizable. The species is
described as having reduced eighth pleurals (which might suggest
a relationship to T. euphraticus), and a narrow mandibular sym
physis without a median ridge. The coloration was described by
Boulenger as blackish green above, elegantly marked with very
numerous roundish yellow spots of unequal sizes; dorsal shield
with interrupted yellow lines, some of which radiate, while others
extend across the costal sutures; plastron mostly greyish.
Trionyx steindachneri is found on Hainan Island and on main
land Kwangtung, Kwangsi, Tongking, and in Viet Nam. It reaches
an altitude of at least 5000 feet in Tongking. It is a small species,
649
www.Ebook777.com
adult at ten inches or less. It is easily identified by the dense patch
of tubercles at the base of the neck; also the mandibular symphy
sis is shorter than the diameter of the orbit, the dorsal tubercles
are arranged similarly to those of T. sinensis tuberculatus, and the
hind extremities of the boomerang-shaped fused epiplastra are
pointed (broad and sometimes dentate in sinensis). The coloration
of the type specimen, a juvenile only about 2.5 inches long, was
given as follows:
“Head olive green above, with only a very few black
dots; a black stripe extends forward from the edge of the eye
to the base of the snout, another obliquely backward to the
neck; likewise two short oblique stripes lie below the eye.
Thus the side of the head is divided into several oval areas,
which are set off by the dark framing as large, lighter spots.
Above the black stripe behind the eye on each side of the
head is a large white spot, which extends backward towards
the neck and turns into a broad band of the same color. The
two spots are separated on the crown by the black color of
the neck, which forms a narrow bridge between them. The
latter divided in front into two crooked stripes, of which the
left runs into a fine line between the eyes, while the right is
severed from the junction. A large, oval, white spot lies just
behind the angle of the mouth. Head and neck dirty white be
low with some darker spots. Carapace dark olive with some
what lighter spots on its margin; a number of diverse, large,
black spots are distributed over the surface. Underside
white with numerous dark, cloudy spots. Limbs uniformly
brown above, somewhat lighter below. ”
The habits of Trionyx steindachneri have not been studied, and
the species is virtually unknown in captivity. It is a highly popular
food item and on Hainan sells for much more than a T. sinensis of
similar size. Pope has remarked that the difference in edibility is
very possibly reflected in the life histories of the turtles, steindach
neri presumably being more vegetarian than sinensis.
GENUS CHITRA
The remaining two Asiatic soft-shells are both gigantic forms
which, because of their peculiar skulls, are classified in separate
genera. The narrow-headed soft-shell turtle, Chitra indica, is
found in the Ganges, Sutlej, and Indus Rivers in India, Pakistan,
and Nepal, and the Ratburi River of Thailand. It probably also
occurs in the Irrawaddy and has been reported from the foot of
Gunong Tahan in the Malay Peninsula. It is a very large species;
there are good records of specimens measuring about three feet in
carapace length and less certain records of much larger speci
mens; Annandale mentions individuals reaching a length of six
650
feet, at which size they are said to be capable of attacking and

sinking small boats. Chitra indica is characterized by the extreme
ly elongate narrow skull; the postorbital bar is around twice the
diameter of the orbit. The eyes are displaced so far forward that
one might be excused for mistaking the orbits in a prepared skull
for the nasal openings. The carapace is dull olive in color, overlaid
in the young by four indistinct ocelli and an extremely complicated
pattern of wavy dark reticulations. In adults the reticulations be
come a less contrasting dark grey. The upper surfaces of the head
and limbs are the same color as the carapace, with a A -shaped
mark on the midline just in front of the carapace and faint yellow
ish dark-edged stripes on the neck. The undersurfaces are white.
Apart from the peculiar skull, Chitra does not differ in impor
tant details from Trionyx. The preneural bone is absent. There are
four plastral callosities and eight pairs of pleural bones, the last
pair being in contact mesially. The anterior rami of the preplastra
are long and slender and widely separated from each other. The
proboscis is relatively short, about the same length as the orbital
diameter. The limbs are flipper-like and excellently adapted for
rapid swimming. The alveolar surfaces are but slightly expanded,
and the edges of the jaws are very sharp; but, like most soft-shells
The incredibly complicated carapace pattern of Chitra indica unfortunately

becomes indistinct with growth. Photo by Dr. W. Sachsse.
651
www.Ebook777.com
except those of the genus Trionyx itself, the narrow-headed soft-
shell is not considered particularly bad tempered or Inclined to
bite.
The habitat of Chitra embraces those sections of large rivers
where the bottom is sandy; it prefers clear water. In diet it is car
nivorous, feeding on fish, molluscs, etc. It is favored for human
consumption and is sometimes offered for sale in Calcutta and in
Thailand. Its reproductive habits are unknown.
GENUS PELOCHELYS
Pelochelys bibroni has a rather broad head, with the orbits
well forward, nearer to the nasal opening than to the temporal
emargination. This species is widespread in southeast Asia, being
known from southern China (Foochow, Canton, Kwangtung, and
Hainan Island) and Burma southward through Viet Nam, Thai
land, and the Malay Peninsula to many of the Indonesian Islands,
including Borneo, Sumatra, Java, and New Guinea. It is also
known from the Philippines (Luzon Island). Whether this distribu
tion reflects transportation by human agency is unknown, but the
species definitely takes to the sea quite freely, although it is typic
ally an inhabitat of fresh water, being found not only in deep slug
gish rivers but even small inland streams.
Pelochelys bibroni, which is known in the Philippines as an-
tipa, is an extremely large species; a specimen reported by Nelly
de Rooij had a carapace length of 51 inches. A Hainan specimen
measured by Clifford Pope had a carapace length of 22 inches and
weighed 42 pounds, which would suggest that de Rooij’s turtle
would have weighed at least 400 lbs. Pelochelys has no preneural
bone; there are seven neurals and eight pairs of pleurals, the last
pair of which meet on the midline. The anterior rami of the pre-
plastra are rather short and broad and are well separated. The
A large specimen of
Pelochelys bibroni,
widely distributed in
southern Asia.
652
snout is short and rounded, and the proboscis is very short. There
are four plastral callosities. There are also callosity-like patches
of roughened scales on the heels, the posterior surfaces of the hind
limbs, another across the outer part of each forelimb, and three in
the web on the outer side of the forelimb. The carapace is olive
with a few darker and lighter striations along the median dorsal
part. The carapace grades smoothly into the fleshy neck area an
teriorly, and the olive color is continued onto the head. There may
be light areas on the temporal region. The dorsal surfaces may be
speckled with yellow, white, or black. The plastron is uniformly
unpigmented. The carapace of the young is densely speckled with
black.
This species is reported to be sluggish and non-aggressive, al
though irritated specimens may strike with great speed, though
without actually biting. The diet under natural conditions is un
known, but captive specimens will eat dead fish. According to
Rudolf Mell, there may be as many as 27 eggs in a single clutch,
and there may be more than one nesting in a season.
BACKGROUND READING
GENERAL:
Geoffroy St.-Hilaire, E.F. 1809. Sur les tortues molles, nouveau
genre sous le nom de Trionyx et sur la formation des cara
paces. Ann. Mus. Hist. Nat. (Paris), 14, pp. 1-20.
Comp. Zool., 115(6): 163-557.
Mehrtens, J.M. 1966. The captive softshell (Trionychoidae):
Checklist of the world’s softshells and their care. Int. Turtle
and Tortoise Soc. J. 1(1): 14-17, 31-33, 38, 45.
Siebenrock F. 1902. Zur Systematik der Schildkrotenfamilie
Trionychidae Bell, nebst der Beschreibung einer neuen
Cyclanorbis-Art. Sitzungsb. Akad. Wiss. Wien, 111, 1:
807-846.
UNITED STATES SPECIES:
Carr, A.F. 1952. Handbook of turtles. Cornell Univ. Press, Ith
aca, N.Y. 542 pp.
States. Kentucky Univ. Press, 347 pp.
Schwartz, A. 1956. The relationships and nomenclature of the
soft-shelled turtles (Genus Trionyx) of the southeastern
United States. Charleston Museum Leaflet 26: 1-21.
653
www.Ebook777.com
Chapter 8
MONOTYPIC TURTLE FAMILIES
Four living kinds of turtle are so unlike all other extant species
that each is placed in a family of its own. One of these four, the
leatherback sea turtle (Dermochelys coriacea) has such a weird
shell structure that it is sometimes even placed in a suborder
(Athecae) which separates it from all other turtles (Thecophora),
but modern opinion, noting the similarity of the skull structure and
nesting habits to those of other sea turtles, takes a more moderate
view, placing this species only in a family of its own, the Dermo-
chelyidae. The leatherback is logically discussed in the chapter
on sea turtles rather than here.
The other three species are the New Guinea plateless river
turtle, Carettochelys insculpta; the Central American river turtle,
Dermatemys mawi\ and the big-headed turtle, Platystemon meg-
acephalum. Each of these is the type and only species of its family
(respectively the Carettochelidae, the Dermatemyidae, and the
Platysternidae).
FAMILY CARETTOCHELYIDAE
The Carettochelyidae were once a widespread group, early
Tertiary fossils referable to this family having been found in
Europe, North America, and southern Asia. However, the single
living species, Carettochelys insculpta, is restricted to the Fly
River area (including the Strickland, Morehead, Lorentz, and
Stekwa Rivers and Lake Jamur) of southern New Guinea and, ac
cording to a recent report by H. G. Cogger, to the Daly River, a
hundred miles southwest of Darwin in the Northern Territory of
Australia.
Carettochelys insculpta is a fairly large turtle, adult examples
being about eighteen inches in carapace length and up to thirty-
five pounds in weight; the record shell length is 19.5”. The dorsal
surfaces of the animal are grey, while the undersurfaces are whi
tish. The jaws are cream-colored, and there is a prominent white
blotch behind each eye. There are also several rectangular white
areas along each side of the carapace near the margin. The snout
terminates in a blunt, thickened ‘breathing tube’ covered with
very tender, wrinkled skin. The dorsal surface of the tail is
covered with a single line of crescent-shaped scales, 14 to 16 in
656
Detail of the tail of

the Fly River turtle,
Carettochelys in
sculpta.The row of
scales is only one of
many distinctive
characters of this
turtle.
number, which decrease in size from the base to the tip. Prominent
folds of skin extend laterally on each side from the undersurface of
the tail across the thigh region and down the hind limbs.
Carettochelys shows a number of striking parallelisms with
sea turtles; the limbs are paddle-shaped, two-clawed, and, while
not as long or as stiff as the flippers of sea turtles, are much more
adapted for a natatory existence than those of other freshwater
turtles. The head can be pulled back inside the shell, but the neck is
so thick that Boulenger, from a study of preserved specimens only,
assumed that the head could not be retracted. The carapace is
relatively deep and globular, with a median keel toward the rear.
There are no scutes, the shell being covered instead with a layer of
continuous skin. In this regard Carettochelys resembles the soft-
shelled turtles (Trionychidae), but many details of shell structure
differentiate the two families. For example, the peripheral bones
of Carettochelys are complete and well-developed, so there is no
flexible shell margin, and the bony plastron is in no way reduced
but forms a continuous plate without even a median fontanelle. In
fact all of the shell bones are exceedingly thick and solid, and the
entire shell surface, with the exception of the undersides of the
front three and hind three peripherals on each side and the under
side of the pygal bone, is strongly rugose—hence the specific name
‘insculpta.’ This rugosity even extends to the dorsal surface of the
skull.
657
www.Ebook777.com
The hatchling of Carettochelys is about 2*4” in carapace
length and differs from the adults in having a depressed, white-
edged carapace with strongly serrate, saw-toothed sides and a
wavy hind margin. The hatchlings also have a tuberculate median
keel; a poorly-defined and transient polygonal outline around each
of these tubercles may or may not be homologous to the scute
seams of other turtles. The median keel loses its tuberculate char
acter as the turtle grows, and with maturity it becomes exsponged
entirely except on the posterior quarter of the carapace.
Several of the plastral elements of Carettochelys are not rigid
ly ossified together, but rather have cartilaginous connections
which allow a certain amount of flexibility. For example, the front
lobe of the plastron (composed of the entoplastron and the paired
epiplastra) is attached to the hyoplastra by a flexible connection;
the only other chelonians with a hinge immediately behind the
entoplastron are the tortoise Pyxis and the musk turtle Stauro-
typus. There are, moreover, longitudinal lines of flexibility along
the midline (except for the front lobe) and along each bridge, so
that, like sea turtles but unlike almost all other turtles, the chest
can pump in and out slightly as the animal breathes. As with the
sea turtles, there is no bony separation of the obturator foramina
of the pelvis.
Carettochelys swimming. The resemblance in general form to a sea turtle is

obvious. Photo by G. Marcuse.
The neural bones of Carettochelys are somewhat reduced; the

actual pattern of reduction is variable, but typically the seventh
and eighth pairs of pleurals have a complete median connection,
and each remaining pair of pleurals is either separated only par
tially by a kite-shaped neural bone situated anteriorly or else com
pletely separated by two neural bones. The pleural bones them
selves do not become rigidly sutured with the peripherals even in
large adults—another parallelism with certain of the sea turtles.
The peripheral bones of Carettochelys are only twenty in number.
Zoogeographically, Carettochelys is something of an enigma,
since its territory is normally considered the exclusive preserve of
side-necked turtles of the family Chelidae. Indeed, Gadow and
Deraniyagala considered Carettochelys to be a Pleurodire, al
though the genus has none of the normal pleurodire char
acteristics. In fact, if one looks at the structure of the cervical ver
tebrae, Carettochelys is almost exaggeratedly cryptodire.
Ginglymoidy, or lateral doubling of the articular facets of the cer
vical vertebrae, is a peculiarly cryptodirous character, being
associated with stabilizing the neck against lateral flexion when it
is forcibly retracted; Carettochelys not only has these double joints
between the sixth and seventh and between the seventh and eight
cervical vertebrae, but it is also the only turtle with a double joint
between the eighth cervical and the first dorsal vertebra.
Actually, Carettochelys is not unique in being a cryptodire in
the New Guinea/Australia region, but it is significant that the only
other cryptodires known from this part of the world are the true
marine turtles and the giant soft-shell Pelochelys, which is known
to be capable of making prolonged marine excursions. It is likely
that Carettochelys too is far from helpless at sea, and that the
North Australian population reached its present location in com
paratively recent times by this means.
Carettochelys is often described as one of the rarest turtles in
the world; but in fact its range is simply so remote and poorly in
vestigated that it is not possible to make such statements just be
cause travellers and collectors have seen relatively few speci
mens. Brongersma, in fact, reported that the species is 'fairly
common’ in the rivers of the south coast of New Guinea and in
Lake Jamoer, or Jamur, a large body cf freshwater also inhabited
by sharks and sawfish. Nevertheless there are very few specimens
in captivity. Two immature specimens were received by the Bronx
Zoo around 1959, and although one died within a fairly short time,
the other is still alive at the time of writing. The late Prof. Dr.
Robert Mertens had another living specimen in his laboratory in
Frankfurt, Germany, and there are two in the Taronga Zoo in
Sydney, Australia. The London Zoo received three adult specimens
a few years ago, but was forced to sell them due to shortage of
659
www.Ebook777.com
space. Two were transferred to the Bronx Zoo and the third to the
present writer’s collection, but all died within a few months from
stomatitis caused by a Proteus infection combined with acute
osteomyelitis.
Judging by the small amount of growth recorded over many
years by the immature specimens in the Bronx Zoo and in Dr. Mer
ten s'collection, Carettochelys appears to be an extremely slow-
growing animal.
The habits of Carettochelys have not been studied in the wild.
The first Australian specimen on record, caught in the Daly River
by a sports fisherman in 1969, was reported to have been taken in
freshwater with a slow current and a sand and pebble bottom.
Shortly after capture this individual excreted traces of Ficus and
of the freshwater snail Notopala (family Viviparidae). Caretto
chelys has very strong jaws, and the supraoccipital process at the
back of the skull, to which the jaw-closing muscles attach, is not
only very massive but also has huge lateral flanges. The lower jaw
is somewhat scoop-shaped in front and has a slightly ridged, ex
panded surface toward the rear on each side. The upper jaw is un
hooked; toward the rear it too has a broadened, longitudinally
ridged surface on each side.
The eggs of Carettochelys have been figured by de Rooij and
are known to be hard-shelled, spherical, and approximately 1.7” in
diameter. Clutches range from 17 to 27 eggs. The nesting process
has not been described; it would be most interesting to know if
Carettochelys, the only non-marine turtle with flippers, excavates
its nest in an essentially sea-turtle or in an essentially freshwater-
turtle fashion.
An amusing folk-story regarding Carettochelys was related to
the traveller A.J. Jicar at the beginning of this century; the trans
lation of the story was found by the present writer in the box con
taining a Carettochelys skeleton which Jicar presented to the Brit
ish Museum in 1902. The story was related to Jicar by Aniwa, Chief
of Mabudamu on Kiwai Island, Fly River, New Guinea, and the
translation is as follows:
“The freshwater turtle called Minowa, the Caretto
chelys insculpta, first came from a long way up the Fly
River, where the inland tribes used to catch and eat it. No-
one but an inland nation would think of eating this creature
as it was from them that the Kiwai natives first learned the
act of coition, and the subsequent conception of women and
birth of children. The first time a Minowa was seen was a
long time ago, on a mud bank at Demeramuba. A man and
his wife were going past there on their way to their garden,
when they saw two Minowas lying side by side in the mud.
(This turtle is always said to lie on the mud when copulat-
660
ting). The woman said ‘Could not we do this thing that the
strange turtle does?’ The man replied ‘I do not know any
thing of it’. The woman persisted, and on going to their gar
den they laid side by side, as they had seen the turtles, and
after a time, the act of coition took place, and in due course
the woman conceived and a child was bom to them. Previ
ously to this the Kiwai tribe did not know from what source
children came. The men had—in this idea—always lived on
Kiwai Island, and wives were procured by capture from the
mainland tribes. The only use they had for wives was to
work in gardens and catch crabs. To this day a Kiwai native
maintains that the act of connection takes place in a garden
in the open air. The great greed of the Kiwai folk for flesh
foods, and the influence of foreigners, is fast overcoming the
dislike of the natives for killing this turtle, but they will not
do so at present, unless forced to (as they believe that to do
so endangered their chance of becoming a father if a young
man, and his sons impotent, if he happens to have any).”
FAMILY DERMATEMYIDAE
The Central American river turtle, Dermatemys mawi, is the
sole survivor of a primitve and once widespread family, the
Dermatemyidae, which apparently had its earliest beginnings in
Asia in the Lower Cretaceous, and which spread in the early Ter
tiary to North and Central America, Europe, and Africa. The
single living species is found in Belize, north and east-central
Guatemala, and the States of Veracruz, Tabasco, Campeche, Chia
pas, and the Territory of Quintana Roo in southern Mexico; the
northwestern limit of the range is reached in central Veracruz. It
is a large species, both sexes frequently reaching a carapace
length of eighteen inches and, according to Miguel Alvarez del
Toro, individuals sixty centimeters (about two feet) or more in
length are common in the Lacantun and Usumacinta Rivers in
Chiapas.
The vernacular name of this species is tortuga blanca in Vera
cruz, but it is not certain whether this name refers to the white
meat of the animal or to its pale undersurface. In Belize it is known
as hickety, a name which, together with the Spanish form jicotea,
is used for many different turtle species in many different parts of
the New World. For example, in Chiapas the local Pseudemys are
called jicotea, while Dermatemys is called simply tortuga.
Dermatemys has a smooth, somewhat flattened carapace
completely without keels in the adult although a median ridge is
661
www.Ebook777.com
present in the young. The hind margin of the carapace is some
what flared and is serrated in juveniles. The plastron is relatively
large, rounded in front and notched behind, and the bridge is wider
than the length of either the front or the hind plastral lobe. The dor
sal surfaces of the animal are grey or grey-brown in color, while
the plastron is cream-colored and the undersides of the soft parts
are white or pale grey. The shell is relatively thick and heavy, and
in large adults the various shell elements become completely
fused so that the sutures are not visible even in dried bony shells.
The first five pairs of pleurals are separated by elongate neural
bones, hexagonal in shape with the short sides located anteriorly.
The sixth neural may be of normal size or it may be reduced so
that the sixth pleurals meet behind it; the seventh neural is re
duced or absent, and the eighth pleurals may or may not be par
tially separated mesially by a small suprapygal situated anterior
to a large, roughly triangular suprapygal that is always present.
As with many turtle species, the vertebral scutes are relative
ly broad in the young but relatively narrow in the adult. The gular
662
Lateral and plastral views of the Central American river turtle, Dermatemys
mawi. Photo above by J. Alan Holman.
663
www.Ebook777.com
scute is frequently single or only partially split, and there may be a
small azygous scute behind it, but the most striking characteristic
of the scute arrangement of this species is that a row of inframar
ginals, usually four but sometimes five in number, completely sep
arates the ventral side of the marginals from the mid-plastral
scutes on each side. The scutes of both carapace and plastron are
very thin and almost membranous in texture; in large adults the
scute boundaries become obscure and may disappear altogether.
The scutes are also extremely liable to abrasion and mechanical
damage; in contact with rough surfaces, such as concrete, the ani
mal literally wears itself away to the bone in a short space of time.
Such an abraded animal, if placed in deep water away from con
tact with rough surfaces, will heal up but never regains its former
smooth appearance; it will eventually slough off sheets of dead
bone which leave a healed but pitted area beneath.
The head of Dermatemys is relatively small, with a strongly
projecting snout and very large, prominent nostrils. There are no
barbels on the chin. Adult males have a golden-yellow triangular
patch extending over the entire upper surface of the head, with
vermiculations of the same color on the sides. In females and im-
matures the head is dingy grey with obscure vermiculations,
though in juveniles there is a pale line extending backward from
the eye. The jaw surfaces are strongly denticulate with a complex
pattern of projections analogous to the teeth of other vertebrates.
The cervical vertebrae show double (procoelous) joints at each
end of the seventh cervical and biconvexity at the second. Both
fore and hind feet are very large and fully webbed. There is a row
of enlarged scales along the outer surface of the front limbs. The
tail is thick and conical and larger in the male than in the female;
it has two rows of conical tubercles above and one more row on
each side.
Among living turtles, Dermatemys is most closely related to
Staurotypus, and the two genera, together with Claudius, were at
one time placed in the same family. Staurotypus differs from Der
matemys in having a larger head with flat, crushing-type jaw
plates, a strongly tricarinate carapace, and in having a reduced
plastron with narrow bridges and a line of flexure behind the ento-
plastron and epiplastra. Also, the inframarginals are reduced to
two on each side (the axillaries and the inguinals), and the pos
terior neural bone is usually in contact with the first supracaudal
so that no pair of pleural bones has a median suture. Staurotypus
also has eliminated the gular and humeral scutes of the plastron.
The fossil genus Xenochelys, from the Oligocene of North Ameri
ca, appears to be somewhat intermediate between the two genera.
In view of the extreme delicacy of the thin shell scutes of Der
matemys, it is not surprising that this is one of the most aquatic of
664
all turtles; it does not climb up on logs and river banks to bask as
do other turtles, but instead floats passively, often asleep, on the
surface of the great rivers of tropical Mexico and northern Central
America. Wilfred Neill obtained a shell of this species covered
with barnacles in Belize, which suggests that in part of its range at
least it may be found in salt water. In diet Dermatemys is entirely
vegetarian, feeding upon aquatic vegetation in the wild, while cap
tive specimens do well on a diet of apple and banana. Captive juve
niles accept meat and fish, which suggests that they are more car
nivorous than the adults in the wild. Dermatemys is of mild dispo
sition and rarely attempts to bite, and even though it may flail its
limbs with considerable force when picked up, its efforts on land
are so poorly coordinated that many specimens have the greatest
difficulty in walking, while some seem reluctant even to lift their
heads off the ground. On the other hand, in water Dermatemys
swims rapidly and well.
In view of the maladaptation of this species for terrestrial
locomotion, the question of how it nests is a fascinating one for
which we must thank Miguel Alvarez del Toro for the answer. Der
matemys in Chiapas nests from September to November, which
corresponds to the time of greatest rainfall, when the rivers in
which this turtle lives swell considerably. The animal allows itself
to be carried by the floods into shallow rivulets, where it makes its
nest and buries its eggs in the zone between the mud and the decay
ing vegetation at the edge of the water. The eggs are typically
about twenty in number and average about2.3” inlength by 1.3” in
thickness; the shells are very thick and hard. Robert C. Lee found
eggs ready for deposition in February and March in females from
Lake Izabal, Guatemala.
Dermatemys can maintain long periods of activity without
rising to the surface of the water to breathe, and it is probable that
this species can extract much of the oxygen it needs from the
water; when submerged in slightly cloudy water it can be seen
that water is continually taken in through the mouth and expelled
through the nose.
The meat of Dermatemys is universally considered excellent
for human consumption, and in view of both the quality and quan
tity of meat that is obtainable from a single turtle, it is not surpris
ing that the species is persecuted and sells for a high price
wherever it occurs. Indeed, considering the ease with which float
ing sleeping specimens may be captured by a cautious canoeist,
some concern should be expressed for its future.
According to Alvarez del Toro, the otter preys upon Derma
temys with some regularity in Chiapas, and it is a common occur
rence to find empty shells of Dermatemys killed by otters on the
beaches of the large rivers.
665
www.Ebook777.com
FAMILY PLATYSTERNIDAE
The third monotypic family, the Platystemidae, is poorly rep
resented in the fossil record, although one fossil genus (Scutemys,
from the Cretaceous of Southeast Asia) is referred to this family
by Romer. The living species, Platystemon megacephalum, is
fairly widely distributed in Southeast Asia, having been reported
from southern Burma, Thailand, Laos, Cambodia, Viet Nam, and
southern China, including Hainan Island. This is a relatively small
species, large adults having a carapace length of only six or seven
inches. However, the head is massive (often almost half the width
of the carapace) and the tail is about as long as the shell. This spe
cies is peculiar in having a completely roofed-over skull, a condi
tion otherwise found only among the sea turtles and in Pseudemy-
dura and Peltocephalus. Whether this is a secondary re-develop
ment in Platystemon or whether the lineage has had a roofed-over
skull throughout is hard to say. The head is not retractile within
the shell, but is protected not only by its bony top but also by a con
tinuous tough homy scute, bigger than any of the carapace scutes,
that covers the entire top of the head as well as the sides right
down to the comers of the mouth. The jaws are very strong and
hooked, and the stout jaw plates are so extensive that there is only
a narrow band of soft skin extending from the eye to the nostril;
posteriorly the upper rhamphotheca abuts directly onto the sides
of the head plate.
The carapace of Platystemon is somewhat indented anterior

ly; without this indentation it would be impossible for the massive
head with its short neck to be raised above the horizontal position.
The nuchal scute is foreshortened by this indentation and is much
wider than long. The carapace is olive-brown in color and depress
ed, often actually concave in the middle, especially in adult ex
amples. The plastron is flat and quite large, though the bridges are
very narrow. There are three inframarginal scutes in each bridge
which completely separate the plastral scutes from the marginals.
Anteriorly, the plastron is abruptly truncated, the gular scutes be
ing exceedingly wide, while posteriorly the plastron is
emarginated at an obtuse angle. The long tail is armored with
large flat scales which are arranged in only three rows posteriorly
(one dorsal and two ventral) but several anteriorly. The base of the
tail and the thigh region are equipped with homy projecting
tubercles. The vent is situated more posteriorly in the male than in
the female. Musk glands are present in the soft skin near the
667
www.Ebook777.com
bridge. The limbs show no special modifications, although the
forearms are well armored with enlarged flattened scales. The feet
are small and the toes only partially webbed.
Two subspecies of the big-headed turtle are well established:
P.m. megacephalum from southern China (from Kwangsi and
Kwangtung northeast along the coast to Foochow), and P. m. pegu
ense from southern Burma, Thailand, and Indo-China. The Hainan
population, geographically intermediate between these two distri
butions, ’s listed as megacephalum by Wermuth and Mertens, but
according to the illustrations provided by Clifford Pope in his Rep
tiles of China, it would in fact appear to be referable to peguense.
P.m. peguense may be recognized by the presence of dark sym
metrical markings on the plastron, which are typically located on
the posterior edges of the humeral, pectoral, abdominal, and fem
oral scutes and along the midline on the femoral and anal region.
These dark markings are also found around the vent and part way
along the tail and on the outer surfaces of the limbs. Otherwise the
undersurfaces of the animal are virtually unpigmented. P.m. meg
acephalum, on the other hand, has uniformly yellow-brown under
surfaces with no black pigmentation. P.m. peguense also has a
more pointed snout and a broader, more keeled carapace which
often shows interrupted lateral keels and concentric and radial
sculpturing on the dorsal scutes. The head too is differently
marked in the two races, peguense having unmarked pale yellow
jaws, a strong, pale, black-bordered line extending horizontally
from the eye to the back of the head, and only a slight amount of
pigmentation on the top of the head. The head of megacephalum is
dark above and below, with numerous orange spots behind and
below the eye and on the lower jaw; the postocular stripe is usually
absent.
A third subspecies, Platystemon megacephalum vogeli, has
recently been described from Chiang Mai Province in northwes
tern Thailand. This race is reported to be distinguished from meg
acephalum by the more narrowed snout and the presence of a dark
figure on the plastron, and from peguense by the shorter, less
beaked upper jaw, by the smooth carapace lacking distinct growth
annuli, the smooth, non-scalloped hind margin of the plastron, and
the greater extent of the dark plastral figure. I am inclined to ques
tion the distinctness of this form. It was based upon only two speci
mens (a living male and a preserved female), and the compara
tive material consisted of a live megacephalum which had been in
captivity for several decades and a live peguense which too had
done most of its growth in captivity. The observed differences
probably result from the unnatural conditions of captive life
(which, for example, can cause formation of unnaturally clear
growth annuli and extreme scalloping of the carapacial margin) of
668
Two views of the big-headed turtle from Southeast Asia, Platysternon

megacephalum peguense. Photos by G. Marcuse.
www.Ebook777.com
the specimens of the other two subspecies, while the different ex-1
tent of the dark plastral figure would have to be confirmed by the
examination of very large numbers of specimens from known lo
calities before it could be considered a convincing difference.
The bony structure of the shell of Platysternon shows few
peculiarities; there are two suprapygal bones, and the neurals ex
tend in an unbroken series from the nuchal bone to the anterior
supracaudal. The nuchal bone is devoid of costiform processes.
The plastron is not rigidly sutured to the carapace, but the two are
only connected by cartilage, so that there is some free movement
at this location in life. The cervical vertebrae show biconvexity at
the fourth vertebra, while the seventh is biconcave with double
joints at each end; the joint between the fifth and sixth vertebrae
may also be doubled.
The hatchlings and young of Platysternon have a stronger
medial keel than the adults, and the posterior margin of the shell is
serrated. The plastron is pink in young peguense. The young are
capable of emitting a high-pitched squeak when molested. The
habits of this turtle have not been extensively studied in the wild,
but the animal lives principally in fast-flowing streams at high al
titudes and is known to be an unusually skillful climber, reportedly
ascending trees and rocks in search of food and for basking pur
poses. The large head and strongly depressed shell move the cen
ter of gravity of the animal well forward, while the reduced
bridges allow the forelimbs to be thrust a long way back, so the
animal is capable of climbing over (or out of) anything which it
can reach with its outstretched claws. When handled Platysternon
can deliver painful bites and holds on with tightly clenched jaws.
Pope reported that specimens hiss when disturbed and turn
toward the captor with mouth gaping. The diet under natural con
ditions has not been studied, but captive specimens eat meat,
snails, worms, and fish. Only two eggs are normally laid at a time.
Platysternon is eaten in Burma and China and is often sold in
the market in Kowloon.
BACKGROUND READING
CARETTOCHEL YS:
Brongersma, L.D. 1958. The animal world of Netherlands New
Guinea. J.B. Wolters, Groningen; 70 pp.
Gogger, H.G. 1970. First record of the pitted-shelled turtle,
Carettochelys insculpta, from Australia. Search 1:41, 1 pi.
de Rooij, N. 1915. Reptiles of the Indo-Australian Archipelago.
I. Leiden.
670
Peters, U. 1970. Die Papua-Schildkrote (Carettochelys insculp-

ta) in Australien! Die Aquarien und Terrarien Zeitschrift,
23(6): 182-183.
Schmidt-Westrum, T. 1963. Die Papuaschildkrote aus Neu-
guinea. Natur und Museum, Frankfurt am Main, 93: 119-127.
Schodde, R., I. Mason and T.O. Wolfe, 1972. Further records of
the pitted-shelled turtle (Carettochelys insculpta) from Aus
tralia. Trans. R. Soc. S. Aust., 96(2): 115-117.
Walther, W.G. 1922. Die Neu-Guinea-Schildkrote Carettoche
lys insculpta Ramsay. Nova Guinea, 13 (Zool.): 607-704.
Zangerl, R. 1959. Rudimentare Carapaxbeschuppung bei jun-
gen Exemplaren von Carettochelys und ihre morphogenetis-
che Bedeutung. Festschr. Steiner. Vierteljahrschr. Natur-
forsch. ges. Zurich 104, 138-147.
DERMATEMYS MAWI:
Alvarez del Toro, M. 1960. Los Reptiles de Chiapas. Inst. Zool.,
Tuxtla Gutierrez, Chiapas, Mexico, 204 pp.
Lee, Robert C. 1969. Observing the tortuga blanca. Int. Turt.
Tort. Soc. J. 3(3): 32-34.
PLATYSTERNON MEGACEPHALUM:
Mertens, R. 1960. Schwimmunfahige Wasserschildkroten.
Natur and Volk 90(4): 127-133.
tral Asia, Vol. 10.
Smith, M.A. 1931. Fauna of British India, Vol. 1: Loricata, Tes-
tudines. London, Taylor and Francis; pp. i-xxviii, 1-179.
Wermuth, H. 1969. Eine neue Grosskopfschildkrote, Platyster
non megacephalum vogeli, n. ssp. Die Aquarien und Ter
rarien Zeitschrift, 22(12): 372-374.
__________ 1970. Eine Grosskopfschildkrote als Kletterkunstler.
Die Aquarien und Terrarien Zeitschrift, 23(1): 23-25.
Wermuth, H. and R. Mertens. 1961. Schildkroten, Krokodile,
Bruckenechsen. Gustav Fischer, Jena.
671
www.Ebook777.com
Chapter 9
SEA TURTLES
Although the earliest turtles in the Upper Triassic and Juras
sic were apparently marsh-dwellers, the move to the marine envi
ronment was a relatively early one in the history of turtles. By the
Cretaceous, marine turtles of various degrees of specialization,
and referable to four families, were already of world-wide distri
bution. Present-day sea turtles are limited to two families, the
Cheloniidae with six species and the Dermochelyidae with a single
species. The Dermochelyidae is an aberrant family of scuteless
turtles which is considered in detail at the end of this chapter; the
discussion here therefore refers largely to the Cheloniidae.
Modern sea turtles have several very primitive features. Con
spicuous among these are the non-retractile head and limbs, com
pletely roofed-over skull, and complete series of inframarginal
scutes. It has been supposed by some that the complete skull roof
is a secondary feature, but there is no evidence for this theory,
which would assume a very rapid and unlikely emargination and
re-formation of the complete skull roof of the ancestral turtles.
This protection for the top of the head is necessary since retraction
is impossible and the marine environment is full of large preda
tors. When attacked while swimming, a sea turtle is able to protect
its flippers to some extent by folding them under the plastron. The
non-retractile extremities in turn are a correlation of the necessity
of streamlining; the anterior carapace overhang necessary in a
turtle with retractile head and limbs would constitute a serious im
pediment to sustained rapid swimming. The flippers of sea turtles
are highly adapted for swimming and are of very minimal efficien
cy on land. The claws are reduced in number (to one or two), and
the long digits are fused throughout their length into coherent
paddle-like structures. The one sea turtle species whose extremi
ties are slightly retractile (the hawksbill) is also almost certainly
the most sedentary species, least in need of streamlining.
Although a sea turtle will occasionally use an alternating paddling
motion in shallow water, the normal mode of swimming involves
powerful simultaneous winglike beats of the foreflippers, with the'
hind flippers assuming a stabilizing and directional rather than
propulsive role. The different terrestrial gaits of living sea turtles
are discussed under the individual species headings.
672
The cervical vertebrae of sea turtles are short and thick; typi
cally the fourth cervical centrum is biconvex. The joint between
the sixth and seventh cervicals is virtually or completely immov
able, the surfaces usually being flat and joined together by strong
cartilage. However, in those cases where this joint is slightly pro-
coelous rather than flat, some lateral doubling (ginglymoidy) may
be present. Were it not for this occasional doubling, a strong case
could be presented for inclusion of the sea turtles in the Amphiche-
lydia rather than the Cryptodira.
673
www.Ebook777.com
Although only seven species of the turtles survive in the oceans
of the world today, the group is a successful and significant one
whose small number of living species is more a reflection of unifor
mity and continuity of the marine environment than evidence of
limited success for the group. The sea turtles are a dominant life-
form in littoral and coastal ecosystems in many parts of the
tropics, though their importance in the economies of both nature
and man has diminished greatly in the last few centuries as more
and more nesting colonies have been exterminated or decimated.
Modem sea turtles have become behaviorally and dietarily
specialized into different niches that provide a minimum of direct
competition; the green turtle is a tropical herbivore, the logger-
head is a subtropical feeder upon heavy molluscs and crabs, the
olive ridley is a tropical species which feeds largely upon relative
ly small crustaceans, Kemp’s ridley is a tropical to subtropical
species which eats organisms intermediate in size between those
consumed by the loggerhead and the olive ridley, the hawksbill is a
tropical reef species which mainly eats sponges, and the leather
back is a tropical to temperate species specialized for a diet of
jellyfish and tunicates. The diet of the Australian flatback is still
poorly known, but even so it appears likely that there is rather lit
tle food competition between the living sea turtle species. In pri
mordial times, however, it is likely that some competition for nest
ing space took place between certain species, and this may still
occur in a few places today. Such competition is minimized by dif
ferences in beach preferences; for example, leatherbacks prefer
Head of a sea turtle, Eretmochelys imbricata, showing the major head scales.
Art by Karen Harrod.
long, muddy, rock-free beaches, while hawksbills often nest in tiny

sandy coves flanked by rocky outcrops. There is also a tendency
for smaller (and thus more shallow-nesting) species to nest later
in the season than larger species when they share the same nesting
beaches. This obviously minimizes the risk of nest destruction by
later turtles nesting in the same spot.
Many aspects of sea turtle classification are still under debate,
but authorities agree that modern species represent two families,
the Dermochelyidae and the Cheloniidae. Some place both these
families in the Cryptodira; others consider the leatherback (Der-
mochelys) to warrant separation at the superfamily level, while
still others place it apart from all other turtles in a separate sub
order, the ‘Athecae’ (or ‘shell-less ones,’ a slight misnomer). Che-
loniid turtles are grouped into four genera; arguments for various
subfamilial placements of these genera are given in the section on
the hawksbill.
Morphologically, modem cheloniids show fewer specializa
tions than some of the late Mesozoic and early Tertiary forms.
Even in the specialization of great size, modern sea turtles are
easily outranked by certain Cretaceous sea turtles of the family
Protostegidae, such as Archelon, which reached a total length of
about ten feet. Still, none of the modem sea turtles are small; the
smallest are the ridleys of the genus Lepidochelys, which are
mature when a little more than two feet in carapace length. The
largest is Dermochelys, whose carapace length measures up to six
feet.
In the phylogeny of most vertebrate lines, a reduction in the
number of bony elements characterizes increasing modernity.
Among the cheloniid genera, it appears likely that increasing
modernity and specialization are marked by an increase in the
number of scales and to a lesser degree in the number of shell
bones. The genus Chelonia may for these purposes be considered
the archetypal, and most primitive, modem form. Eretmochelys
shows a division of the prefrontal scales; Caretta shows further
division of the head scales as well as insertion of an extra costal
scute at the anterior end of each costal series; Lepidochelys not
only shows the division of the head scales and the extra costals, but
also shows extensive division of the neural bones and often division
of the costal scutes to a total of as many as nine on each side. Lepi
dochelys also shows four pairs of inframarginals and the most ex
tensive development of Rathke’s glands known among turtles.
The bony armor of sea turtles is most complete in the ridleys;
the hawksbill turtle, whose scutes are very thick and provide good
protection from abrasion and impact, has the least ossified cara
pace, with the lateral fontanelles between the open rib-ends re
maining open throughout life. In all sea turtles there is some longi-
675
www.Ebook777.com
tudinal flexibility at the bridges and along the midline of the plas
tron; the chest can thus move in and out slightly as the turtle
breathes.
The jaw surfaces of sea turtles show adaptations to the differ
ent dietary specializations of the various species; the vegetarian
green turtle has finely serrated jaw surfaces, which in some popu
lations form quite long pseudo-teeth. The loggerhead and the rid-
leys have jaws adapted for the functions of crushing and grinding.
The hawksbill has a narrow head with the jaws meeting at a highly
acute angle, which facilitates extraction of food organisms from
crevices in coral reefs. The leatherback has delicate scissor-like
jaws which would suffer severe damage if subjected to anything
much harder than the usual food of jellyfish.
In recent years, the concentration of female sea turtles in cer
tain favored areas for nesting and the dispersal of the turtles at the
end of the nesting season have prompted wide-scale tagging pro
grams in an effort to elucidate the migrations of the various spe
cies. Although other kinds of tag have been used occasionally, the
universally favored type is a Monel metal cattle ear tag which is
clipped through the trailing edge of the front flipper by means of a
specially designed pair of pliers. In areas where turtles are not
completely protected by law, these tags offer a reward to the cap-
tor of the turtle, and in this way a picture of the migration patterns
of the turtles is gradually built up. Such results are discussed for
each species later in this chapter; however, it appears that the
green turtle is the most purposeful migrant, making journeys be
tween well-defined and sometimes quite circumscribed nesting
and feeding grounds. The one-way journey may on occasion be as
much as 1,500 miles. The ridieys and the leatherback appear to dis
perse widely after nesting and to re-concentrate for the next nest
ing season; leatherbacks have been recovered over 3,000 miles
from the point of tagging. Loggerheads, which are typically tem-
perate-zone nesters, show a tendency to move into warmer, more
tropical waters during the winter months. Long-distance migra
tions have not yet been demonstrated for the flatback or the
hawksbill. The mechanisms of sea turtle navigation have been the
subject of intensive experimentation and speculation, but we still
do not know what cues or sensory systems are involved.
GENUS CHELONIA
The green turtle, Chelonia mydas, may well be considered the
‘archetypal’ marine turtle; it is the largest of the hard-shelled spe
cies, it is found in all the tropical oceans, and it is of such economic
importance that Archie Carr, the world authority of the species,
has called it ‘the most valuable reptile in the world.’ When Euro
peans first penetrated to the tropical seas, it was in many cases the
676
green turtle that kept them alive; these big, succulent reptiles
abounded in many areas and could be captured without any equip
ment or skill simply by turning them on their backs when they
emerged from the sea to lay their eggs. At the present time the
green turtle has lost its importance as a victualling source for
ships, but it still has great economic importance, not only as a pro
tein source for subsistence-level coastal people, but also as the
only source of ‘real turtle soup’ beloved by high-living Europeans
and the source of most of the turtle steaks which are in ever-
increasing demand, more by novelty-seekers than gourmets, in
resort restaurants in the Caribbean, Florida, Hawaii, and other
places. In view of this enormous demand for green turtle products,
it is not surprising that whole breeding populations have been
wiped out within historical times and many others have been deci
mated; indeed it is amazing that the species has managed to sur
vive at all in some places where exploitation has been intensive for
generations.
The green turtle is a handsome, streamlined creature with a
smooth, heart-shaped shell and a rather small, anteriorly rounded
head. The flippers are long, and the normal mode of progression
both in the water and on land involves simultaneous sweeps of the
forelimbs.
The carapace of the green turtle is dark blue-black in the
hatchling, and this coloration may persist for several weeks or
months. However, before long the central part of each dorsal scute
lightens to a mid-brown color, and by the time a carapace length of
about one foot has been reached, the central and costal scutes have
assumed a pattern of radiating streaks of buff, brown, and black.
The intensity of this pattern is extremely variable even for turtles
of the same size and from the same population; in some individ
uals light colors predominate, in others dark. The dorsal colora
tion of the mature green turtle shows even greater variability; the
background color may be green, olive, brown, buff, or black; radi
ating streaks, either entire or broken, may be present, or the juve
nile pattern may break down completely, reduced either to irregu
lar spots or to a plain overall slate-grey color. In some areas (e.g.
South Yemen) it has been observed that most of the turtles with
the spotted pattern are males, but this does not hold up in all other
areas, and it is possible that it is simply the oldest turtles that be
come spotted and that, in areas where females are killed on the
nesting beaches, nearly all of the old animals are males.
The underside of Chelonia mydas is pure white in hatchlings
with the exception of a band of dark pigment along the underside of
the marginal scutes. This band varies in intensity from one popula
tion to another; it may be intensely black in juvenile Ascension
turtles. The proximal parts of the flippers are white, but the distal
R77
www.Ebook777.com
parts are black. The coloration of the head and flippers tends to be
an extension of that of the shell. However, the edges of the head
scales are pale throughout life, and the soft skin between the scales
of the flippers is light-colored in juvenile specimens.
The plastron remains virtually unpigmented throughout life in
green turtles from most parts of the world, merely becoming pale
yellow instead of pure white in adults. However, in the East
Pacific area the plastron soon acquires extensive areas of grey.
Details of this population, which is considered to be referable to a
distinct subspecies (agassizi), are given below.
The squamation of the green turtle normally consists of four
pairs of costal scutes, five central scutes, eleven pairs of margi
nals, a pair of supracaudals, and a single broad nuchal. The scutes
are rather thin and are juxtaposed throughout life. The plastron in
cludes the usual paired elements as well as one or two intergulars
and sometimes a narrow interpectoral scute. There are four pairs
of inframarginals. The commonest variant dorsal scutation in
volves the inclusion of a small extra central scute between the
fourth and fifth large scutes. Occasionally there may be an extra
costal scute on one or both sides. The nuchal scute, unlike that
of the ridleys and the loggerhead, does not make contact with the
first costal scute on each side. The head scalation typically in
cludes four postocular scutes and is normally symmetrical, with
out the extensive scale fragmentation shown by Caretta and Lepi-
dochelys. The largest scale on the crown of the head is the fronto
parietal, which is preceded by a smaller frontal and a single pair
of elongate prefrontals. On each side of the frontal is a supraocu
lar, and behind each of these is a temporal. A pair of parietals is
situated behind the frontoparietal, with an additional temporal on
the outer margin of each parietal.
The head of the green turtle is relatively small, with large
orbits, more or less parallel sides, and a short, rounded snout. The
alveolar surfaces are rather complicated, there being strong
ridges parallel to the outer edge in both upper and lower jaws; the
lower jaw has a strong ‘tooth’ at the rear of the symphysis, and this
fits between and behind two similar though blunter ‘teeth’ in the
upper tomium. The outer margin of the lower jaw surface is den
ticulate, especially in certain East Pacific populations, and these
denticulations correspond to vertical grooves in the inner surface
of the cutting edge of the upper jaw.
The adult size of the green turtle differs from one part of the
world to another. A typical mature green turtle of either sex has a
carapace length of about 40 inches and a weight of 300 to 350 lbs.
East Pacific green turtles, which correspond to the subspecies
agassizi, are smaller, averaging about 32” at maturity (smallest
about 28”; largest about 39”). On the other hand, certain areas, in
678
cluding Surinam and Ascension Island, are characterized by green

turtles of enormous size. The average carapace length in Surinam
is about 44”, while the largest specimen known from there had a
48” carapace and weighed 493 lbs. In 1973, I found a green turtle in
French Guiana measuring 50.5” (over the curve). In Ascension
the maximum size is still greater; the largest green turtle
measured anywhere in recent years was a 55 ” female from Ascen
sion. The heaviest green turtle caught in modern times was a
pathologically obese sterile female from Grand Cayman Island;
although only 49” long, it weighed 649 lbs. Its shell is still to be seen
in Ira Thompson’s Museum on Grand Cayman. The biggest speci
men ever caught is supposed to have weighed 850 lbs.
At the present time, nesting of the green turtle in the continen
tal United States is limited to a handful of females which come
ashore each summer on the Atlantic coast of Florida. In former
times, however, nesting was much more common in Florida, and
immature green turtles were common even off North Carolina,
where they are virtually unknown today. Moreover, a century ago
green turtles of enormous size (up to 1000 pounds according to
True, though this weight estimate is usually discounted nowadays)
were common near Cedar Key, on the Gulf Coast of Florida. Today
mature greens are rarely seen north of the Florida Keys, and the
few still found at Cedar Key usually weigh fifty pounds or less.* In
the Caribbean there are still two major nesting sites for the green
turtle: the beach that extends south for 22 miles from Tortuguero,
on the Caribbean Coast of Costa Rica; and the half-mile sandbank
known as Aves Island, in the middle of the East Caribbean, over
130 miles from the nearest land of any sort. On a peak-season
night, two or three hundred turtles may nest along the entire beach
at Tortuguero and two or three dozen at Aves. The Tortuguero
rookery has been studied more intensively than any other; Dr.
Archie Carr, of the University of Florida, has maintained a re
search camp there every summer since 1955, and a tremendous
volume of data on the nesting population has been accumulated.
Tortuguero was also the source for baby turtles for ‘Operation
Green Turtle,’ a program designed by Dr. Carr, and supported by
the U.S. Navy, for taking newly hatched green turtles to various
Caribbean beaches from which the nesting population had been
extirpated, in an attempt to imprint the babies on these beaches so
that when they reached maturity they might return there to nest.
Unfortunately no new breeding colonies appear to have been es
tablished, and a few years ago the experiment was discontinued.
*Nevertheless, some believe that the green turtle is making a

comeback in Florida, and summer nesting emergences on Hutch
inson Island and Merritt Island are becoming almost routine.
679
www.Ebook777.com
Hatchling green turtles, Chelonia mydas , from Costa Rica released in Bimini.
The Aves Island rookery is not nearly so well known, since the
logistics of even getting to the island, let alone working there for
protracted periods, are very complicated. However, William
Rainey of the Island Resources Foundation on St. Thomas has now
begun a tagging program there.
Elsewhere in the Caribbean, some greens nest on the short
Caribbean coast of Mexico, especially on Isla Mujeres and the
coast of Quintana Roo. Green turtles are still found in some num
bers on the south coast of Cuba and presumably still nest there, but
the Grand Cayman green turtles were exterminated long ago. On
the coast of South America, the best nesting grounds are in Suri
nam (Bigisanti, Baboonsanti, Eilanti, and Pruimenboem
beaches), though nesting also occurs on Shell Beach in Guyana
and on Silebache Beach, French Guiana. The status of the green
turtle on the coast of Brazil, at least as a nesting animal, is largely
unknown, but some nesting supposedly takes place on Marajo
Island and near Rio Doce and Cabo Frio. There is still a populous
680
colony on Trindade Island. The most important mid-Atlantic

colony is that on Ascension Island, where the turtles have not been
commercially exploited for many years and are thus still very
plentiful. Nesting also occurs in the Cape Verde Islands. No data
are currently available on the status of the green turtle as a
nesting animal in West Africa. A few are found in the eastern Med
iterranean, and nesting is known to occur at Yamortalik on the
coast of Turkey.
In the Indian Ocean, the most important green turtle nesting
grounds are on the mainland coasts of the Arabian Peninsula,
especially on the coast of South Yemen but also on the east coast of
Saudi Arabia; green turtles are reported to nest on numerous
islands in the Persian Gulf and on Masira Island. Nesting is also
known to occur in Somalia, Tanzania, and Mocambique, but nest
ing in Kenya is now sporadic. There is a major nesting beach at
Hawke’s Bay, Karachi, and nesting is also known in the Comores,
Maldives, Diamond Island (Burma), and on Cocos-Keeling Island.
The green turtle still nests in good numbers on both coasts of Thai
land and Malaya, especially on the offshore islands. Green turtles
are now greatly depleted in the Seychelles and the various BIOT
islands; perhaps the best remaining nesting colonies are on Astove
and Cosmoledo Islands. The once great rookery on Aldabra is now
almost gone, but the colony on tiny Europa Island in the Mocam
bique Channel, having been completely protected since the 1920’s,
may now have the largest green turtle colony in the world; George
Hughes, who had himself stranded on this island for several weeks
in November 1970, was surprised to find about 700 turtles nesting
every night on only about four miles of beach. Although obviously
pleased by his discovery, he was also somewhat frustrated, having
only brought some 400 tags.
Australia has some green turtle nesting grounds of great im
portance, the best-known being certain coral cays in the Capricorn
and Bunker groups at the southern end of the Great Barrier Reef,
especially Heron Island; other important nesting areas are Raine
Island and Bramble Cay at the northern end of the Great Barrier
Reef, and Bountiful and associated islands in the Gulf of Carpen
taria. Some nesting almost certainly occurs in Western Australia,
but this is not yet documented. In Indonesia the green turtle is
known from the islands of Sumatra, Biliton, Borneo, Djawa,
Madura, Flores, Sulawesi, Ambon, Banda, Ori, Aru, and Irian. In
Borneo nesting occurs not only in the Indonesian part of the island,
but also on many beaches in Sabah, especially on the islands of
Pulau Selingaan, Pulau Bakungaan Kechil, and Pulau Gulisan.
During the 1970 season, about 750 green turtles are estimated to
have nested on the best of these beaches, that of Pulau Selingaan.
The three famous turtle islands of Sarawak have substantial,
681
www.Ebook777.com
though dwindling, populations of green turtles; about half of the
eggs are laid on the island of Talang Talang Besar, about 40% on
Talang Talang Kechil, and about 10% on Satang Besar. In the old
days the numbers of eggs harvested in these islands were astro
nomical; more than 3,000,000 were collected in 1936. However, at
the present time, the Sarawak turtle population is in serious
trouble, suffering from well over 90% exploitation of the eggs laid
as well as increasing destruction of the adult turtles by fishermen
from the Philippines and elsewhere. The averge annual egg yield
from 1927-1936 was 2,147,000; from 1967-1972 it was 321,000.
The seven Philippine turtle islands, in the Sulu Sea quite close
to Sabah, together constitute an important green turtle nesting
ground. In 1953, nearly 4000 eggs were collected nightly on one
island, Taganak, and nearly as many on Baguan.
A complete enumeration of the Pacific Islands on which green
turtles nest is beyond the scope of this book. However, the species
still nests in reduced numbers on French Frigate Shoal, in the
Hawaiian Leewards, and has the unusual habit of basking on the
beach in the heat of the day on this and on other Hawaiian Islands,
including Pearl and Hermes Reefs and Lisianski and Laysan
Atolls. Diurnal emergence of female turtles for protracted periods
has also been observed on Momington Island in the Gulf of Carpen
taria, Australia. The local Lardil aborigines consider such turtles
come ashore to escape the attentions of over-zealous males, and
Robert Bustard, who has visited the island, concurs with this
opinion.
In March and April, 1976, I conducted a preliminary study of
the marine turtles of the U.S. Trust Territory of Micronesia. Nest
ing occurred in several of the districts; in Palau the most frequen
ted are the uninhabited or sparsely inhabited southern islands of
Merir and Helen’s Reef, while in the Yap District green turtles
nest on Ulithi (islets of Gielap and Iar), Ngulu, Pikelot, West Fayu,
Ifaluk, and Olimarao. In Truk District some greens nest on East
Fayu, and on Oroluk in Ponape District. In the Marshall Islands
the best nesting is on Bikar and Jemo, but some greens nest on
Taka, Erikub, Ujelang, Jaluit, Taongi, and Bikini. In most areas
nesting takes place in the summer months (May to July), but on
certain islands, notably East Fayu and Ngulu, some nesting
occurs as early as February.
Harold Hirth, after a survey of green turtle stocks in the South
Pacific, concluded that the most important breeding areas today
were: Fanuatapu, Namu’a, Nu’ntele, and Nu’ulua in Western
Samoa; Rose Atoll in American Samoa; Scilly, Mopelia, and Bel-
linghausen Islands in French Polynesia; the Ha’apai Group in
Tonga; Lau, Yasawa, and Mamanutha Groups in Fiji; and lie
Ouen, lie Surprise, and Recif d’Entrecasteaux in New Caledonia.
682
In the East Pacific, green turtles are found from the central part of
the Gulf of California south to central Peru. In Mexico, nesting
takes place in the Revillagigedo Islands, in considerable numbers
near Maruata Bay in the State of Michoacan, and apparently in
even larger numbers in Oaxaca, about 25 km. west of Salina Cruz.*
Nesting also takes place at Playa Azul, Michoacan, and in front of
Mar Muerto, Chiapas. Major nesting grounds for this species in
Central America have not yet been found, but good numbers nest
on certain of the Galapagos Islands. The best nesting beaches in
the archipelago are situated on the north coast of Santa Cruz
Island (Indefatigable); on the west coast of Baltra Island (South
Seymour); on the north, east, and west coasts of Santiago Island
(James); on the south coast of Bartholomew Island; on the north
east coast of San Cristobal Island (Chatham); on the north coast of
Floreana Island (Charles) near Cormorant Point; and on Bara-
hona and Quinta Playa on the south coast of Isabela Island (Albe
marle). In the Galapagos Islands, in addition to the normal green
turtles (locally known as “black turtles”), there exists a minority
of individuals, mostly females, which are distinguished by their
bright yellow plastra, thick carapace shields, and tremendous
obesity. These so-called “yellow turtles,” greatly sought-after in
former times for their high oil content, are never seen on the nest
ing beaches and never contain eggs at any time of the year. While
the situation here merits close study, my tentative explanation is
that these “yellow turtles” are a sterile mutant form of the “black
turtle.” Tremendously fat barren green turtles are also occasion
ally found in the Caribbean—the 649 pound female from Grand
Cayman mentioned above was one of them.
A recent report in Science by Felger, Cliffton, and Regal con
firmed the amazing phenomenon of winter dormancy in green tur
tles in the north-central Gulf of California. In this area, during the
cold months (approximately November to March), the turtles
repair to certain habitual areas where they shuffle into the bottom
mud so that they are almost concealed; they remain torpid until
the onset of spring. Such hibernacula are now known near the
islands of Raza, Angel de la Guarda, Salsipuedes, San Lorenzo,
San Esteban, San Pedro Martir, and Tiburon.
Since about 1959, the Seri Indians of the coast of Sonora have
exploited these turtles commercially, catching them by low tide on
clear, calm days when the water is relatively clear and the turtles
may be spotted by the exposed central part of the carapace show-
*The Maruata turtles nest from June to October; six or seven times
during the season arribadas of about 1000 turtles come ashore,
although the ‘background’ nesting level is nearer 50 turtles or
fewer each night.
683
www.Ebook777.com
ing above the substrate. The turtles are harpooned and, being very
sluggish, are easily brought ashore.
The commercial exploitation of these dormant turtles in
creased enormously in 1972-73 when Mexican fishermen in Kino
Bay discovered them. Since then, several of the principal bedding-
down sites have been denuded of turtles by these fishermen, who
operate with professional diving equipment and gasoline-powered
air compressors which allow them to remain submerged for one or
two hours at a time. It appears that these unique winter-dormant
turtles will be exterminated within a year or two of their being
brought to the attention of the scientific world.
Although the green turtle has been subject to intensive study
for many years now, the various subspecies that surely exist have
never been properly defined. The only local form whose validity is
universally accepted is the East Pacific green turtle, Chelonia my
das agassizi, which occurs in the Galapagos Islands and also on the
mainland Pacific coasts of the Americas. This race is character
ized by its tendency toward very dark dorsal pigmentation coupled
with extensive areas of grey on the plastron. The carapace shape
also differs from that of C.m. mydas in usually being higher and
more steep-sided, often with distinct indentations in each side of
the carapace above the hind flippers. The size is also smaller—the
average weight of mature females is well under 200 pounds. Else
where in the world, the populations of Ascension Island, the
Guianas, and Aves Island are distinguished by their great size and
by subtle differences in shell shape, but these have not yet
received trinomial recognition. The amazing thing about the re
maining populations—for example those of Costa Rica, the Sey
chelles, the Great Barrier Reef, and the Sarawak Turtle Islands—
is how very similar they are; what differences may exist have yet
to be elucidated.
Tagging experiments in several parts of the world have dem
onstrated that green turtles may nest several times in a season, at
an average interval of two weeks between nestings. The Sarawak
records accumulated for many years by Tom Harrisson show that
six or seven nestings in a season are not uncommon, while a few
turtles nested 8, 9, 10, or even 11 times. Miss Kay Pau, of Damley
Island, Torres Strait, recently observed (and indeed assisted—the
turtle had a mutilated flipper and could not dig its own nest cavity)
a green turtle nesting eight times on Damley Island; eggs were
counted on each occasion, and successive nests contained 154,166,
170, 172, 159, 161, 140, and 98 eggs, for a season’s total of 1220. The
small final clutch seems to be fairly typical for the species. On the
other hand, Archie Carr is becoming increasingly convinced that
some green turtles lay only one clutch in a season at Tortuguero,
Costa Rica, and Johan Schulz believes that the average green tur-
684
tle at Bigisanti, Surinam, may lay only 1.5 times per season,
though some lay five or six times.
Nesting by green turtles in successive years is very rare; typi
cally the majority of individuals nest every third year, with a
minority (though often a significant one) nesting every two years.
Individuals have been known to switch from a three year to a two
year cycle, but it is not known whether it is simply older turtles
that nest at shorter intervals, or whether it is better feeding condi
tions, or higher sea temperatures, that sometimes allow a turtle to
get back into breeding condition after only two years. It is quite
possible that some turtles nest on a four-year cycle, but this is diffi
cult to prove without an enormous volume of statistical data and
very good coverage of the nesting beaches, since it is always pos
sible that a turtle seen again four years later re-nested after only
two years but was not seen. Throughout the world, the interval be
tween nestings within a season appears to be of the order of twelve
days to two weeks. This interval, incidentally, is also typical of
hawksbills, loggerheads, and flatbacks, though leatherbacks have
a rather shorter average interval and ridleys are very irregular.
The nesting process of the green turtle does not vary signifi
cantly from one part of the world to another, and, with minor vari
ations described later in this chapter, the nesting of this species is
typical of sea turtles generally. A green turtle emerging from the
sea to nest, however, is more sensitive to disturbance than is a sea
turtle of any other species. But the degree of skittishness even
among green turtles varies; in Sarawak it is reported that even the
light from a cigarette half a mile down the beach will scare them
back. In the Guianas the greens are not that sensitive, but I well
remember several years ago hiding behind a log on the beach at
Bigisanti, Surinam, to take a flash photograph of a green turtle as
she ascended the beach. As soon as the flash fired, I heard a rapid
slapping as the turtle ran back to the sea, and by the time my eyes
had re-accustomed themselves to the darkness, the turtle was
nowhere to be seen. On the other hand, the ridleys and leather
backs nesting on the same beach could be photographed and even
tagged as they left the sea without frightening them back.
Nesting of the green turtle is seasonal in some parts of the
range, virtually year-round (though sometimes with seasonal
peaks) in others. Nesting is almost always by night, though on
some beaches (e.g. in the Galapagos Islands) the first few turtles
may emerge before the sun has disappeared. The turtle usually
moves very slowly as she hauls herself laboriously up the beach,
and this phase is punctuated by frequent, complete stops. When the
turtle has finally reached the dry sand area out of reach of the
highest tides, she usually does little further wandering before com
mencing the excavation of her so-called “body pit.” The body pit,
685
www.Ebook777.com
which when completed lowers the whole turtle 9 to 18 inches below
the undisturbed sand level, is initially dug with simultaneous sand
sweeping movements of the two fore flippers, which are brought
forward, palms outward, until they almost touch, then thrust
strongly backward through the sand. Before long one of the hind
flippers also starts kicking, then the other, and finally kicking by
the hind flippers alternates between periods of sweeping of the
fore flippers. The movement of the fore flippers gradually dies
down, and the hind flipper movement, from being a general sand
throwing, becomes a specific effort to dig out the sand immediate
ly under the area of the tail. This digging movement, which is car
ried out by alternate actions of each hind flipper, eventually re
sults in the formation of the actual egg cavity. Each flipper is in
serted in the cavity, whereupon it makes several scraping move
ments at the front wall of the hole, then the loosened sand is with
drawn with the deftness of the human hand and gently deposited
beside the cavity. The action is then repeated with the other flip
per, and just before the first flipper is re-inserted in the hole, it
flicks sharply forward, thus preventing a buildup of sand beside
the hole. During this time the eyes of the turtle are emitting
copious viscous “tears.” These tears not only serve to eliminate
excess salt from the system, but also to keep the eye free of sand
when the beast is on a beach and throwing sand in all directions.
When the egg cavity is completed, the hind flippers are dis
cretely folded over the tail and oviposition commences. The eggs
are dropped in batches of between one and about four and when
laid are surrounded by thick clear mucus. The number of eggs laid
varies from one population to another; in many areas the average
clutch contains between 100 and 110 eggs, but in Surinam the aver
age is 142, while in the Galapagos Islands the average is between
70 and 90, very few nests containing over 100 eggs. The average
egg diameter ranges from about 1.6” in Sarawak to about 2.2” in
Ascension, though in certain areas (including Hawke’s Bay in
Pakistan and the Galapagos Islands) each clutch may include a
few markedly undersized, yolkless eggs.
When all the eggs are laid, the turtle sweeps sand into the cavi
ty with alternate movements of the hind flippers; a mound of sand
soon forms, which the turtle pounds and kneads for some time.
Later the front flippers start moving again, throwing sand back
ward with a movement similar to that used in the initial excava
tion of the body pit. The sand-throwing movements continue, and
eventually the turtle starts inching forward with each stroke. This
barely perceptible motion finally brings the turtle two or three
yards beyond the spot where the eggs were laid; the body pit has
now been moved forward with the turtle, and the eggs are exceed
ingly hard to find at this point without the use of a sharp ‘probe’
686
stick. Eventually the turtle starts a deliberate humping-forward

movement which carries her back to the sea, after two hours or
more on land.
On many nesting beaches nowadays, relatively few green tur
tle nests remain intact. Man is a widespread and efficient predator
upon the eggs in many parts of the world, and indeed in some
places, particularly in Southeast Asia, local people have been col
lecting such a high proportion of the eggs for such a long time that
it is a miracle that the turtle populations still survive. Animal
predators upon the eggs include raccoons, feral dogs, monitor liz
ards, and ghost crabs. The hatchlings in an undisturbed nest
appear at the surface about two months after the eggs were laid,
having spent the last two or three days climbing through a foot or
more of sand. The turtles are known to work together as they
scratch at the ceiling of the egg cavity, gradually raising the floor
with the loosened sand until they get very close to the surface. Dur
ing the daytime, when surface sand temperatures are above about
29° C., the turtles are inhibited by the heat and will not make the
final breakthrough to the surface until the cool of the evening (or
until the surface sand is cooled by rainfall). The final eruption is
‘explosive,’ with perhaps a hundred baby turtles breaking out
simultaneously and immediately rushing for the sea. Their orien
tation is usually good even when the sea is out of sight, though
sometimes a hatchling will wander inland a good deal before final
ly getting its bearings, and thus run an even higher risk than usual
of being eaten by a predatory bird or crab. Massive mortality
awaits them in the ocean too, but turtles have evolved for millions
of years in the face of such predation, and if only the breeding fe
males are left alone, the species will survive. Incidentally, it is not
unknown for the adult turtles as well as the hatchlings to lose their
orientation on land, particularly when they are nesting on beaches
with high dunes and with no high land or forest inland from the
beach. In such situations the normal cue of greater light intensity
from the sea side may be absent, and some of the turtles will
wander inland until they become exhausted, whereupon they will
die in the sun the next day. This happens on Bikini Atoll in the Mar
shall Islands, as well as on Baltra Island in the Galapagos.
The green turtle is unique among sea turtles in having a
largely (though not entirely) herbivorous diet from the age of
about one year onward. The main food is made up of sea grasses in
some areas, though in others various species of algae make up the
bulk of the diet. The former diet is said to be more conducive to the
formation of meat of superior quality for eating purposes. Known
genera of food plants for Chelonia mydas include Zostera, Sagit-
taria, Vallisneria, Thalassia, Cymodocea, Deplanthera, Gelidium,
Sargassum, Rhodymenia, Gracilaria, Griffitsia, Liagora, Ulva,
687
www.Ebook777.com
Gratelonpia, Caulerpa, Halodule, Halophila, Syringodijm, Ptero-
cladia, and Posidonia. Jellyfish are also known to be eaten, as well
as various molluscs, crustaceans, echinoderms, and sponges but
these form only a small fraction of the diet in most areas. The
plants which make up the bulk of the diet may form submarine
pastures of enormous extent, and it is on these pastures that the
turtles graze for years at a time. However, it so happens that the
best nesting beaches are often hundreds of miles from the best
feeding grounds, and accordingly green turtles have evolved
strongly migratory habits. For example, it is now known that
green turtles that nest on Ascension Island feed on the coast of
Brazil, at least 1400 miles away. The turtles from Surinam and
French Guiana, curiously enough, feed in the same area. Thus,
when the breeding season approaches, some of the Brazilian tur
tles will migrate eastward across hundreds of miles of open ocean
to a small island breeding ground, while others will set off in the
opposite direction, swimming along the coast for many hundreds
of miles in a westerly direction to the beaches of the Guianas. The
Tortuguero green turtles are mainly derived from feeding grounds
in the Miskito Cays, a few hundred miles to the north; but others
come from the coasts of Panama and northwestern South Amer
ica, and there have been occasional recoveries even from the
Yucatan Peninsula, Cuba, and Belize. Comparable patterns of
migration are emerging for green turtle populations in other parts
of the world, though one requires the combination of a very large
breeding population together with extensive fishing on the feeding
grounds before sufficient data can accumulate to give a quantita
tive picture of where the turtles go.
The mechanisms of migration and navigation in the green tur
tle have been studied intensively by Dr. Archie Carr for many
years. The various possible hypotheses are reviewed in detail by
Dr. Carr in his book So Excellent a Fishe. Suffice it to say here that
we still do not have the final answer, and indeed the feat of locating
Ascension Island, barely seven miles across, after negotiating
1,400 miles of open ocean, by an animal that is hopelessly myopic
when it raises its eyes above the level of the water, seems little
short of miraculous.
Even very young green turtles may make extensive migra
tions. A 20-ounce captive-raised specimen released near Delray
Beach, Florida, on April 13, 1972 was recaptured on May 6,1973, at
Aves Island off the coast of Venezuela, 1300 miles away as the crow
flies and a little further by the shortest sea route. During this
period the turtle increased its weight to nearly four pounds.
The male turtles migrate to the nesting grounds with the fe
males, and during the early part of the nesting season copulating
pairs may be seen floating together for hours at a time. The male
688
turtles, which are immediately recognizable by their long tails,

have recurved claws on the foreflippers which allow them to grip
the front edge of the carapace of the female in the shoulder region.
Female turtles often bear deep notches in the shell in this area,
and the notches appear bloody and raw shortly after copulation. In
many parts of the world several males may be seen fighting each
other for access to a female; however, such sights probably result
from a biased sex ratio caused by excessive killing of females on
the nesting beaches. On Europa Island, where the population
structure is unaffected by the activities of man, George Hughes
reports that the females may even fight over males! ‘Natural’ sex
ratios are unknown, but appear to approach equality in some
areas, though not in others.
Sex ratios of green turtle samples from eggs laid in the wild
but hatched in captivity have been assembled for various locations
and years by the researchers at the Grand Cayman Island Turtle
Farm. The results are exceedingly interesting. The ratio of males
to females ranged from 437 males to 92 females (i.e. 17.42%
females) for a batch of 1970 hatchlings from Ascension Island, to 1
male to 250 females (i.e. 99.6% females) for 1971 turtles from Suri
nam. The turtles were sexed when slaughtered for the market at
an age of about four years. There was no correlation of sex ratio
with origin—the 1969 Ascension turtles were 82.6% females, yet the
1970 batch was 17.42% females. The most probable explanation,
according to David Owens, an associate researcher at the turtle
farm, is that sex is labile in the embryos, and that it differentiates
as the eggs or hatchlings develop according to the prevailing tem
peratures. Certainly it is hard to improve on this hypothesis, which
has fascinating implications for the population structure of wild
turtles.
I believe I was the first to witness the initial stages of the para
doxical phenomenon of attempted nesting by male green turtles;
in early 1970, while camped on Espumia Beach on James Island in
the Galapagos, I saw a turtle emerging from the surf, and I hid be
hind a bush to watch its progress up the beach. The turtle climbed
all the way to the dry sand area above the high tide mark, then
noticed me and started returning to the sea. It was then that I saw
it had a tail at least a foot long and was obviously a male. What it
would have done if I had not disturbed it is hard to say, but in Suri
nam in both 1972 and 1973, two or three male green turtles seen by
Forest Service Personnel not only made the complete ascent of the
beach, but also excavated deep nest cavities! I can offer no explan
ation for this weird and unusual phenomenon. Two of the males
which voluntarily came ashore in this way in Surinam in 1972
ended up in the breeding tank at Mariculture Ltd. on Grand Cay
man Island. Despite well over a month out of the water after cap-
689
www.Ebook777.com
ture, they were seen copulating within minutes of being placed
with females in the breeding tank.
There has been some debate as to whether copulation off a
nesting beach results in the fertilization of eggs for that season or
for the following season. However, most authorities now believe
that the current season’s eggs are being fertilized, partly because
copulation is observed mainly at the beginning of the nesting sea
son (one would expect it toward the end of the season if it were
the next season’s eggs that were being fertilized), partly because
the alternative hypothesis leaves unanswered the question of when
the virgin females are fertilized.
The ardent passion of a male green turtle is often exploited by
turtle fishermen, who either take advantage of the male’s preoccu
pation to harpoon him from a small boat or else attract him by
means of a crude wooden decoy which may or may not be carved
into an approximate representation of a female turtle. The males
are said to cling to these artifacts with such persistence that they
may be simply reeled in and hauled aboard. Perhaps the most
amazing method of turtle fishing, however, and one which is prac
ticed by completely isolated peoples as far apart as Cuba, Kenya,
and the Torres Strait, Australia, involves the use of the remora,
Echeneis naucrates. This fish is equipped with an oval suction pad
that covers most of the top of the head, and in nature it uses this to
hitch free rides with sharks, turtles, and other large marine ani
mals. The fishermen in the areas mentioned attach a line to a ring
around the tail of the remora and release the fish in an area where
turtles are known to occur. It is said that it does not take long for
the fish to attach itself to a turtle, and when it does so the turtle is
secured simply by pulling on the line attached to the suckerfish
(though two or three fish may be needed to make sure a large tur
tle does not pull free and escape).
Until very recently, it was usual to read in most herpetological
books and papers that the genus Chelonia only included the single
species Chelonia mydas. However, as long ago as 1880 Samuel
Garman described the very distinctive form Chelonia depressa
from northern Australia, and the type specimen, an adult male,
had been available for study in the Museum of Comparative Zoo
logy ever since, while for many years one of the equally distinctive
hatchlings had been in the British Museum. D.B. Fry in 1913 made
a valiant but largely ignored effort to demonstrate the distinctness
of depressa, but it was not until fifty years later that Eric Worrell
re-admitted the validity of the species, and this was done in a
casual and imprecise way which made no mention of the fact that
herpetologists throughout the world had been ignoring the species
for many years and also confused the species with a still indefinite
form of green turtle from Japan, Chelonia japonica (Thunberg
690
1787). However, in 1967, Ernest Williams, Alice Grandison, and

Archie Carr (Breviora, M.C.Z. No. 271) re-examined the literature
and the available specimens and formally re-instated Chelonia
depressa as a valid species.
The flatback turtle is a little smaller than the green turtle, the
average over-curve carapace length for fourteen mature females
being 36.3”. The carapace, as both the vernacular and scientific
names imply, is strikingly depressed, also rather wide, and with
distinctively turned-up sides. The carapace scute counts are the
same as those of Chelonia my das. The scutes are olive-grey in
color in the adult and have a thin, greasy character, with seams
that almost disappear with age. The plastron is uniformly white,
with a relatively high frequency of postanal scutes.
The head of depressa is relatively larger than that of mydas,
and from above shows more of a triangular than a rounded shape.
There are only three postocular scales on each side (green turtles
almost always have four), and the postparietal scales, which num
ber two in mydas, typically number three in depressa, although
some individuals have only one and others show asymmetrical
splits into as many as four. In color the head is greyish buff,
becoming yellowish below.
The front flippers are relatively shorter than those of mydas,
and whereas those of mydas are covered with relatively large
polygonal scales, those of depressa have large scales only on the
edges, most of the area overlying the phalanges being covered
simply with finely wrinkled skin. This reduction of scalation is also
apparent on the hind flippers.
Still other characters for distinguishing mydas and depressa
have been described; these include the uniformly small upper eye
lid scales of depressa, the small number of subtemporal scales
(behind the postoculars), the rare or limited contact of the pre
frontal and the maxillary sheath, the reduced axillary scutes, and
the relative proportions of the plastral scutes, with the interfem-
oral seam being the longest in depressa.
The flatback turtle is at present known definitely only from
Queensland, the Northern Territory, and northern Western Aus
tralia. It is typically a tropical animal, but it is known to nest on
both Curtis and Facing Islands, just south of the tropic of Capri
corn off the Queensland coast, while small numbers nest even
further south on Mon Repos Beach, a mile-long beach frequented
by larger numbers of nesting loggerheads. Mon Repos is at a lati
tude of about 25° S and is the southernmost known nesting record
for the species. Tropical nesting beaches for the flatback turtle in
clude the islands of Indian, Baresand, Quail, and West Peron, near
Darwin, N.T. Crab Island, off extreme northern Queensland, is an
important rookery for the species; Alvar Mould and three assis-
691
www.Ebook777.com
tants tagged 180 flatbacks on this one island in one month in late
1970. Perhaps the most interesting observation made there was
that nearly all nestings were diurnal, the peak hour for emergen
ces being about 3:00 p.m. Nesting has also been recorded in the
Coburg Peninsula area and on the Sir Edward Pellew Islands in
the Gulf of Carpentaria. In Queensland, nesting is known on the
Pisonia and Bountiful Groups, near Momington Island, and fresh
track counts suggest that well over a hundred turtles may nest
each night on each of these islands. The southernmost major
rookery is near Gladstone. Although the species has never been
definitely recorded outside Australia, Deraniyagala recently sug
gested that a turtle he found in Ceylon many years ago may have
been a flatback.
Flatback turtles nesting at Mon Repos have been studied in some
detail by Colin Limpus, of the National Parks and Wildlife Service.
The nesting season, as one would expect, takes place in the south
ern summer, late October to late January. On the other hand, in the
tropical Northern Territory, Worrell reports that nesting takes
place all year round. Limpus was able to ascertain that up to four
clutches may be laid in a season, with recorded inter-nesting inter
vals of 13 to 18 days (mean 15.3 days for a sample of nine). The
delicate skin and carapace of the nesting female only showed
mating scars at the beginning of the season, so it is assumed that,
as with other sea turtles, mating takes place shortly before the first
nesting.
Little is known of the behavior of the flatback turtle, and its
migrations, if any, are completely unknown. The species is
reported to be of a nervous and somewhat pugnacious disposition
while on the nesting beach. The gait is the typical Chelonia syn
chronized action of all four flippers, but the movements on land
are much brisker than those of Chelonia mydas\ one turtle seen
nesting on Crab Island was out of the sea for only about forty
minutes. In the sea flatbacks spend much of the day floating at the
surface, quite frequently with sea birds perched on their backs. In
this respect, they would appear to show a similarity to the olive
ridley of the East Pacific.
One of the most interesting facts about Chelonia depressa is
the small number and large size of its eggs. The flatback is essen
tially a medium-sized sea turtle, mature females averaging about
160 pounds and thus smaller than the green, loggerhead, or
leatherback, and larger than the hawksbill or either of the ridleys.
Yet 87 nests examined by Limpus contained, on average, only 50.2
eggs each (range 7-73). This is far fewer than the average for any
other sea turtle. On the other hand, the eggs average 1.97” in diam
eter and are thus as big as leatherback eggs and distinctly bigger
than those of all other cheloniid sea turtles. One would suspect that
692
this device, which presumably represents a division of the maxi

mum possible egg biomass into a rather smaller number of rather
larger packages than usual, is a means of producing hatchlings
which just exceed the maximum size to be killed by certain poten
tial predators. This indeed appears to be the case. There are, for
example, two ghost crab species on Mon Repos Beach; the smal
ler, Ocypode cordimana, can sometimes drag a loggerhead hatch
ling into its burrow but is not large and strong enough to drag off a
flatback hatchling, which is probably about 50% bigger than a
hatchling loggerhead. The larger crab species, Ocypode ceratoph-
thalma, has no trouble killing a hatchling green turtle or logger-
head, but only occasionally is able to kill a hatchling flatback.
Also, silver gulls, which are able to eat hatchlings of both of the
other two species, appear not to molest flatback hatchlings.
The hatchlings of Chelonia depressa are quite distinct from
those of C. mydas. Apart from being about 40% larger, the cara
pace has a serrated margin and a broadly oval, rather than heart-
shaped, outline. The carapace scutes are yellow with grey mar
gins, while the underside is white with a yellow band along the
periphery. The head is grey and white, and the eyes have a most
attractive turquoise color.
The feeding habits of the flatback turtle have not been proper
ly studied. Worrell reports that the stomachs he examined con
tained brown-colored seaweeds and cuttlefish. There is also some
evidence that flatbacks feed upon trepang, a collective name for
several species of sea cucumber of the genera Actinopyga and
Holothuria. The short flippers and broad skull suggest that this
species may be more of an opportunistic forager than a regular
migrant between well-separated feeding and nesting grounds, as
are most populations of C. mydas. The flesh of the flatback has
been reported to have a disagreeable taste, but Worrell found it
quite palatable.
The survival outlook for the flatback turtle appears to be good,
since many of its nesting beaches are located in uninhabited
aboriginal reserves and steps are being taken to establish national
parks for the more important Queensland nesting beaches. How
ever, both monitor lizards and introduced foxes are serious preda
tors on the nests in many areas.
GENUS CARETTA
In former times the loggerhead turtle, Caretta caretta, was
confused with the hawksbill, Eretmochelys imbricata, and this
confusion is reflected both in the scientific name of the species (de
rived from carey, Spanish for hawksbill), and in the modem Dutch
vernacular name, Onechte Caretschildpad, or false carey turtle.
In fact the two species are quite different, and the loggerhead is
more closely related to the ridleys than to the hawksbill.
693
www.Ebook777.com
The loggerhead is a large sea turtle with a rather elongate,
red-brown carapace. The shell tapers posteriorly, and the widest
point is reached well in front of the mid-point. There are five cen
tral scutes and five pairs of costals. The marginal scutes usually
number eleven or twelve pairs (not counting the paired supracau-
dal scutes). The shell is very thick in an adult loggerhead. The
shell thickening is particularly extreme in the region of the hind
part of the fifth central scute and the supracaudals—in an adult
loggerhead a solid hump of bone is present in this area. The pos
terior thickening of the shell probably lessens the likelihood of dis
abling bites from sharks, from which this rather slow-swimming
turtle has little chance of escaping by flight. The carapace scutes
are rather thin and are juxtaposed throughout life in contrast to
those of the hawksbill, which are thick and imbricate. The hind
margin of the carapace is serrated. The plastron is yellowish and
typically (though by no means invariably) has only three infra
marginal scutes in each side. No inframarginal pores are present.
The carapace of a halfgrown loggerhead has a distinct spine at the
rear of each central scute.
The head of the loggerhead is very large; this is especially
striking in adult specimens, not because they have a relatively lar
ger head than a young turtle, but because they do not show the
usual allometric reduction in relative head size with growth that
almost all animals show. Needless to point out, however, the large
head of an adult loggerhead does not house an enormous brain, but
simply very large jaw-closing muscles. In color the head and limbs
of the loggerhead have the same red-brown coloration as the shell.
The ventral surfaces of the extremities are a dirty yellowish color.
The head scales of the loggerhead are rather complex. A frontal
and large frontoparietal are always present, the former some
times divided. There are two pairs of prefrontals, which in some
cases surround a small azygous extra scale.
Two subspecies of the loggerhead have been described: Caret-
ta c. caretta in the Atlantic and C.c. gigas in the Indian and Pacific
Oceans. The subspecies are supposed to be separable on the basis
of the number of marginal scutes and of the number of neural
bones, as well as on the frequency with which the neural bone
series is interrupted by the pleurals. While it is quite possible that
in due course the loggerheads of the different oceans will be shown
to be distinct, the characters mentioned above do not hold up. For
example, it is stated that caretta has, on average, 11 marginals
(not counting the supracaudals) on each side, while gigas
averages 12. However, the actual average for 43 West Atlantic log
gerheads was 11.62; for 12 Western European specimens it was
11.71; for six West African specimens it was 11.83; for seven Medi
terranean specimens ft was 11.07; for 26 Indo-Pacific loggerheads
694
An adult loggerhead turtle, Caretta caretta. The relatively large size of the
head is obvious, and the head scalation is visible. Photo by G. Marcuse.
(from Australia, the Solomon Islands, and Ceylon) it was 11.78; for
7 Mexican Pacific loggerheads It was 11.07; for 76 Japanese log
gerheads it was 11.44. The marginal count is thus useless for differ
entiating subspecies.
The neural count difference (supposedly 7 or 8 in caretta, 7-12
in gigas) has not yet been systematically disproved; however,
twelve specimens from Ceylon examined by Deraniyagala had an
average neural count of 9.1; eleven specimens from the Atlantic
(from Ireland, Holland, Georgia, Florida, and Colombia) had
exactly the same average, 9.1. We shall therefore use the binomial
Caretta caretta for all loggerheads until the use of trinomials for
the different populations is properly justified.
The loggerhead has what might be called an “antitropical”
breeding range—almost all of the nesting areas, except for those in
the western Caribbean, are situated north of the Tropic of Cancer
or south of the Tropic of Capricorn. Possibly because of similarity
in the feeding habits, the loggerhead seems unable to co-exist with
the olive ridley, and where one of these species is abundant the
695
i
www.Ebook777.com
other is virtually or utterly absent. In the Atlantic the best breed
ing grounds are located on the Atlantic coasts of Florida, Georgia,
and the Carolinas. Some nesting takes place on most beaches with
in this area, but well-defined “rookeries” exist, for example at
Cape Romaine, South Carolina; Little Cumberland Island and
Jekyll Island, Georgia; Hutchinson Island, Juno Beach, Jupiter
Island, and Cape Canaveral, Florida. By far the best county in
Florida for loggerhead nesting is Brevard, followed by Martin and
Palm Beach. Nesting is sparse north of North Carolina, but there
are a few records for Virginia and even Maryland, while in 1972 the
northernmost nesting record for a sea turtle anywhere in the world
was registered at Ocean City, New Jersey. Sparse loggerhead
nesting occurs along much of the shoreline of the Gulf of Mexico;
on the lower Gulf Coast of Florida, fairly good loggerhead nesting
beaches exist on Cape Sable and Keewaydin Island, and smaller
numbers nest on Sanibel Island. On the Gulf Coast of Florida very
little nesting occurs north of Sarasota County, with the exception
of Franklin County, where nests are found regularly.
In the western Caribbean area loggerheads nest on the
western part of Cuba and in small numbers in the Cayman
Islands; in Quintana Roo, Mexico; on various cays (particularly
San Pedro, Ambergris Cay, Northern Two Cayes and Half Moon
Cay, Sapodilla Cayes, and Hunting Cay 1 off the coast of Belize; the
islands of San Andres, Providencia, and the Albuquerque Cays in
the Western Caribbean; the coast of the Santa Marta Peninsula,
Colombia; and (occasionally) on the mainland of Venezuela,
various of the Venezuelan offshore islands, and Trinidad. The spe
cies is caught off Puerto Rico but is not known to nest there. Some
nesting takes place in the Bahamas, and the species is often caught
in Bahamian waters.
The two major discontinuities in loggerhead nesting on the
Atlantic coasts of the Americas are in Central America from
Nicaragua to Panama and in South America fro Trinidad through
eastern Venezuela, Guyana, Surinam, French Guiana, and pro
bably much if not most of the coast of Brazil. However, the log
gerhead is commoner than the olive ridley in Ceara, Brazil, while
the loggerhead is reported to be the commonest sea turtle in
Uruguay; these turtles probably nest in Espirito Santo, Brazil.
In the Mediterranean the loggerhead is known to nest in Tur
key and Israel as well as in Sicily, Sardinia, Italy, and Libya. It is
probable that at least some nesting occurs along the entire Medi
terranean coast of North Africa. It is likely that nesting occurs
along parts of the coast of West Africa, but precise data are lack
ing.
The loggerhead is often found in areas considerably north of
its breeding range. On the American side there are records from
696
as far north as Nova Scotia. There are also records for Bermuda
and the Azores, and the species is apparently very plentiful in
Madeira, several thousand specimens being caught annually for
the manufacture of souvenirs. There are more records of the log
gerhead from the Atlantic coasts of Europe than of any other sea
turtle, including over fifty records for the British Isles. The two
most extreme records are for Loch Lomond, Scotland, in 1861, and
for the Barents Sea, Russia, in 1964; the former is completely land
locked except for the River Leven, while the latter is ice-hound for
most of the year! Loggerheads from northern Europe are fre
quently small in size, and only rarely are mature specimens found
there. The turtles are usually found in the winter months, when
they are benumbed by low temperatures and allow themselves to
be captured or stranded.
In the Indian Ocean, loggerheads nest in Mocambique from
the Paradise Island group near Vilanculos, southward along the
entire coast to the Tongaland coast of Natal. Concentrations ap
parently exist on the seaward shore of Bazanito Island and on the
Cape San Sebastian Peninsula. Loggerheads also nest in the
southern part of the Malagasy Republic, especially in the Fort
Dauphin area, and on the west coast at least as far north as Moron-
dava. Most of the remainder of the Indian Ocean is thoroughly
tropical and is also inhabited by the olive ridley, so little logger-
head nesting takes place. A conspicuous exception, however, is
Masirah Island, off the southeastern part of the Arabian Penin
sula. Harold Hirth estimated that at least 150 loggerheads nested
each night on one 5-km. section of beach on this island. Moreover,
in temperate Western Australia at least some loggerhead nesting
occurs in the Shark’s Bay area, and it is possible that nesting of the
species is widespread in southern Western Australia.
In the Pacific, loggerheads nest in the southern islands of
Japan in moderate numbers; nesting takes place on all coasts of
Kyushu as well as on most of the offshore islets; on the south coast
of Shikoku; and on southwestern and southern Honshu as far north
as Kashima Nada. In the south temperate zone, loggerheads nest
in eastern Australia, especially on certain islands in the
Capricorn-Bunker Group at the south end of the Great Barrier
Reef, including Heron Island and Wreck Island, and at Mon Repos
Beach near Bundaberg, Queensland. Loggerheads have been
found at sea in Tasmania and New Zealand. A few nest in the Solo
mon Islands and also in Fiji.
There are no known loggerhead rookeries in the Central
Pacific, and in the East Pacific, which is the world headquarters
for the olive ridley, the loggerhead is virtually unknown as a nest
ing animal and is only known from a handful of stray individuals
from Mexico. It is probably food competition from the olive ridley
697
www.Ebook777.com
together with the loggerhead’s predilection for subtropical nesting
grounds that prevents colonization of the tropical East Pacific. In
the north and south temperate zones of the East Pacific area it is
likely that the cold North Pacific current and the Humboldt Cur
rent respectively produce cooling of the coastal waters to a level
too low for successful colonization and nesting by the loggerhead.
The maximum size reached by the loggerhead turtle has been
obscured and exaggerated by miscalculations of overall size based
upon the size of the head. For example, Deraniyagala wrote in 1939
that
“there is good reason for supposing that the brown-red
loggerhead of the Indo-Pacific is the largest member of its
family, for many of the skulls collected were scarcely smal
ler than those of Dermochelys.”
In fact the loggerhead has a disproportionately large head, rela
tive to those of other sea turtles. A leatherback with a head nine
inches wide has a carapace length of about 62”, while a loggerhead
with a head of this width has a carapace length of only about 40”.
Carr (1952, Handbook of Turtles) calculates weights of two logger-
heads with skull widths of 10 inches and 11.2 inches respectively as
1,067 and 1,192 pounds. However, he bases these calculations on a
statement by True in 1884 that a loggerhead with an eight inch
wide skull weighed 850 pounds. In fact, a loggerhead with a head of
that size only weighs about 220 lbs. and has a carapace length of
about 36”. Calculations on this basis give the length of the logger-
head with the 11.2” head as 49.8” and a weight of about 600 lbs.,
which is still gigantic but at least believable.
Mature female loggerheads in the United States, South Africa,
and Japan have a mean carapace length of 36” to 38”; maturity
may be reached at as little as 31.25”, while the largest specimen on
definite record had a carapace length of 45.25”. The normal weight
of mature loggerheads in most areas is from 200 to 350 lbs., but in
Australia few exceed 150 pounds. The loggerheads in Colombia
appear to be slightly smaller, on average, than those from temper
ate nesting grounds; the mean carapace length there is about 34”.
Males are of about the same length as females, but have much
longer tails, recurved claws on the flippers, and flatter and nar
rower shells.
The loggerhead is primarily carnivorous in diet; its broad,
massive jaws have tremendous crushing strength. Brongersma
(1972, European Atlantic Turtles) summarizes the diet of the log
gerhead as follows:
“We may state that on the high seas loggerheads feed on
Scyphomedusae, Physalia physalis, Vellela vellela, Salpae,
Pteropods, Ianthina, Nautilograpsus minutus, Lepas anati-
fera, squids (inter alia Leachia), Syngnathid fishes (Ente-
698
lurus aequoreus), and perhaps also on other fishes. When in

shallow, coastal waters bottom dwelling organisms enter
into the diet, such os crabs, hermit crabs, conchs, borers, sea
urchins, and sponges. Apparently algae are sometimes
taken in great quantity, and, when feeding at the surface
sea-weeds will be taken with other food. The turtles will
swallow floating objects, sometimes inadvertently, some
times because goose-barnacles are attached to these ob
jects. ”
The loggerhead, like other sea turtles, is of a relatively peace
able disposition when on the nesting beach, but individuals moles
ted while at sea may make a considerable show of temper and can
be dangerous to handle. A delightful account of a loggerhead beat
ing up five men who tried to kill it and nearly destroying their row
boat appeared in the New York Herald of September 3,1905, and is
quoted in full in Archie Carr’s Handbook of Turtles and in my
Living Turtles of the World. Nevertheless, the loggerhead, when
plucked from the sea, is not quite as outrageously hysterical as
Kemp’s ridley, and loggerheads will at least survive when placed
on their backs on a ship.
The nesting of the loggerhead shows few noteworthy peculiari
ties. The animal has a typical reptilian alternating gait on land.
The ascent of the beach is quite rapid, though there may be one or
more stops for breath and circumspection during the ascent. Once
the high tide mark is passed, the turtle soon stops and “appraises”
the sand both by thrusting its snout into it and by making trial
sand-sweeping movements with the front flippers. Usually the tur
tle is satisfied with the location and will proceed with the excava
tion of the body pit without delay; this behavior contrasts with that
of the green turtle, which is very fussy in its choice of nest site and
often makes lengthy pauses while making up its mind. The body
pit of the loggerhead is rather shallow and is excavated with just a
few minutes of rather disorganized action of all four flippers. The
final pit is deeper posteriorly than anteriorly, but the carapace of
the turtle remains well above the surface of the sand. The excava
tion of the egg cavity follows the rigidly stereotyped action se
quence shown by all sea turtles, though the movement of the cara
pace from side to side between insertions of opposite hind flippers
is more pronounced, for example, than that of the green turtle.
During oviposition the hind flippers are stretched out flat on each
side; their hind margins curl upward as each batch of eggs is
dropped. The eggs cannot be seen at this point, since the carapace
extends rather far posteriorly and completely obscures the egg
cavity. During the process of filling in and disguising the nest area,
the turtle alternates sand-sweeping movements of the front
flippers with periods of pressing sand down with the “knees; ” dur-
699
www.Ebook777.com
ing this time the posterior part of the carapace rocks from side to
side, but the action is nevertheless quite distinct from the shell
rocking movement shown by ridleys at this stage. A nesting log
gerhead is usually out of the sea for about two hours.
In nearly all parts of the range, the loggerhead nests by night;
however, on Heron Island, in the Great Barrier Reef of Australia,
the turtles often nest in the late afternoon, when the high tide hap
pens to occur at this time. The eggs of the loggerhead average
about 1.64” in diameter in the United States, though some are as
small as 1.4” and others almost 2” in diameter. Recorded dimen
sions of loggerhead eggs from South Africa and Colombia are vir
tually the same as these. Average clutch size for 71 South Carolina
nests was 126 (range 64 to 198). However, at Sanibel-Captiva the
recorded mean is only 110, and two samples in Colombia, the for
mer based on 93 nests, gave respective averages of 108 and 105
eggs. In South Africa the average of 98 nests was 112. Incubation
normally takes about two months. A sample of 26 nests in South
Africa showed an overall average fertility of 88.3%, while 82.55%
of the eggs produced emergent hatchlings. The hatchlings have an
average carapace length of about 1.8”. In color they vary from
pale tan to deep red-brown. There are three keels on the carapace
and two on the plastron. The central scutes are relatively much
wider than those of the adults. In some areas it is not uncommon to
find one or two non-viable albinos in a nest; this albinism is often
correlated with mouth deformities and cyclopism.
The fate of a hatchling loggerhead once it reaches the sea is
not well known. However, several recent observations in Florida
and elsewhere suggest that the baby turtles become associated
with mats of floating Sargassum weed; during fall and winter
storms on the Atlantic coast of Florida it is not unusual for baby
loggerheads to be thrown up on the beaches in association with
clumps of Sargassum. The coloration of the hatchling loggerhead,
with red-brown pigment both above and below, is highly cryptic in
the Sargassum environment.
Few data are available on the migrations of adult logger-
heads. However, it is known that Florida loggerheads may travel
some hundreds of miles up or down the coast and probably move
out to the Bahamas in some numbers during the winter months.
One loggerhead, tagged while nesting on the Atlantic coast of
Florida on Jensen Beach, was caught off the mouth of the Missis
sippi about a year later. Bustard recorded three at-sea recoveries
of loggerheads tagged at Heron Island in Australia; one from Bun-
daberg, less than 100 miles to the south; one from Weipa, in the
Gulf of Carpentaria; and one from the Trobriand Islands. Hughes
obtained evidence that loggerheads tagged in Natal, South Africa,
frequently swim north to Mocambique and Tanzania after nesting.
700
The loggerhead nests several times in the course of a season;

present evidence suggests that three or four nestings per season
may be typical and that the average interval between nestings is
from 14 to 17 days. It is probable that normally nesting years alter
nate with non-nesting years, but there are a few cases on record of
loggerheads in Australia and in South Africa nesting in successive
years.
GENUS LEPIDOCHELYS
The ridley turtles of the genus Lepidochelys are clearly closely
related to the loggerhead turtle, Caretta caretta, but nevertheless
there is no doubt that they warrant inclusion in a separate genus.
After centuries of nomenclatorial confusion, which has been sum
marized in my doctoral dissertation, it is now established that the
genus contains two full species, neither of which has recognized
subspecies. Kemp’s ridley (Lepidochelys kempi), also known as
the Gulf ridley, the Mexican ridley, the Atlantic ridley, or the bas
tard turtle, is confined to the Gulf of Mexico and the north Atlantic
Ocean, while the olive ridley (L. olivacea), also known as the
Pacific ridley, is found in coastal waters of the tropical parts of the
Pacific, Indian, and South Atlantic Oceans.
Kemp’s ridley is a small sea turtle; mature females are usual
ly 26 to 27 inches long and are not known to exceed 29.5”. Breeding
size females range from 80 to 110 pounds in weight; mature males
have not been measured in numbers, but are probably of similar
size. The carapace of Kemp’s ridley is flattened and exceedingly
broad—in many immature individuals the shell is as wide as, or
wider than, long, and in adults the width is typically only a couple
of inches less than the length. The head is relatively rather large
and triangular in shape when viewed from above. The orbits are
rather small, and the jaws are massive, with longitudinal ridges
on both the bony and horny surfaces. Hatchlings are dark grey-
black in color, often with traces of white on the umbilical promin
ence and on the plastral ridges. As the turtle grows, the plastron
loses its dark pigment, becoming white in immature specimens
and yellow as maturity is approached. The dorsal surfaces also be
come lighter with growth and in the adult are olive-green in color,
though half-grown specimens are usually grey and very old speci
mens may revert to a dark grey color. The ventral yellow color ex
tends dorsally to the sides of the head and neck and the entire
shoulder region.
The carapace surface is composed of five central scutes, five
pairs of costals, twelve pairs of marginals, a single nuchal, and a
pair of supracaudals. As with all sea turtle species, these meristics
are somewhat variable; for example, it is not uncommon to find
Kemp’s ridleys with a small extra central scute just behind the
fourth central. However, the costal count is impressively constant,
701
www.Ebook777.com
especially in view of its extreme variability in L. olivacea. The
plastron consists of paired gular, humeral, pectoral, abdominal,
femoral, and anal scutes; an interanal may be present, and there
may be one or two intergular scutes. There are nearly always four
pairs of inframarginal scutes, each of which is perforated toward
its hind margin by a pore which leads to a Rathke’s secretory
gland (these glands are found in the axillary and inguinal regions
of many turtles). The shell of Kemp’s ridley is well ossified, with
no open fontanelles between the distal ends of the ribs in mature
specimens. The neural bones are variable in number but very
numerous; in many cases what would correspond to a single
neural bone in other turtles is split into two separate bones in Lepi-
dochelys. The division is usually transverse, but in some cases is
longitudinal. The front flippers of Kemp’s ridley are relatively
short, and each bears only a single visible claw in mature speci
mens, though hatchlings show a very small distally-located second
claw on each fore flipper and there are usually two claws on each
hind flipper.
Apart from the trivial exceptions noted below, Kemp’s ridley
only nests on one beach in the entire world, and largely because
this beach is in a remote part of Mexico (in the southern part of the
State of Tamaulipas near the village of Rancho Nuevo in the Muni-
cipio de Aldama), the reproductive habits of the species were
entirely unknown to the scientific world until 1963. In that year a
16 mm. film made in 1947 by Andres Herrera was shown at the
annual meeting of the American Society of Ichthyologists and Her
petologists. This amazing film showed an estimated 40,000 ridleys
nesting in a massive synchronized diurnal reproductive effort,
locally known as an ‘arribada.’ These ‘arribadas’ were said to take
place three times a year, in the months of May and June, and to
occur without warning on a slightly different stretch of beach each
time. It was not until the mid-1960’s that Mexican conservationists
succeeded in establishing a camp on the nesting beach to afford
protection and study of the nesting turtles, and by this time the
colony (and thus the world population, considering mature fe
males only) had been reduced to 5000 turtles or less. The causes of
this reduction were not well documented, but are almost certainly
associated with intensive egg-collection by the local people, with
considerable destruction of the remaining nests by coyotes, to
gether with extensive slaughter of the turtles on the beach and fre
quent accidental drowning of adult turtles by the trawlers which
work the west coast of the Gulf of Mexico in considerable numbers.
In 1973 the largest arribada numbered only about 1000 turtles, and*
the second largest only about 250, though on various other days
during the season between one and a couple of dozen turtles
nested. Kemp’s ridley must now be considered a seriously threat
702
ened species, and while it is gratifying to know that the Mexicans

are now working exceedingly hard to protect both the turtles and
their eggs on the nesting beach, the harder task of limiting the
activities of the shrimp trawlers lies ahead. However, the species
does now enjoy complete legal protection in both Mexico and the
U.S.A., and the National Marine Fisheries Service is initiating
research in reducing the accidental losses in shrimp trawls.
The nesting habits of the ridley are peculiar in a number of
ways. The first oddity is that the turtle nests by day rather than by
night, as is the case with all other sea turtles almost everywhere.
Perhaps correlated with this diurnal preference is the fact that the
orbit of Kemp’s ridley is relatively much smaller than that of all
other sea turtles (including the olive ridley, which is normally a
nocturnal nester). Moreover, the habit of forming nesting aggre
gations is shared only with certain tropical American populations
of the olive ridley. The survival value of the trait can only be
guessed, but it is often thought that this is a means of swamping
predators with a huge but very temporary over-abundance of food
('food’ may mean the turtles themselves, their eggs, or the hatch
lings). Thus the resident predator populations will be kept within
reasonable limits, since high populations could not be maintained
by only one or two big meals a year, and when the arribada does
take place, some eggs will survive even though the predators
(principally coyotes nowadays, although it is also possible that in
former times the adult turtles were preyed upon by jaguars, which
are still found in the vicinity) will eat all they can throughout the
duration of the manifestation. It is worth noting that the track of
the ridley is extremely shallow, the turtle being broad and very
light, and also that nesting only takes place during relatively high
winds (the one small arribada I saw in 1973 followed a night of
hurricane-force winds of up to 81 mph). Thus, after an arribada the
tracks are quickly obliterated by blown sand, and the odors associ
ated with the nesting turtle and the fresh eggs also disappear
quickly. So a nest that survives the critical first 24 hours is likely to
remain unmolested until it hatches.
How ridleys establish and maintain their aggregations is not
known at the present time, but it is possible that the secretion of
the unusually well-developed Rathke’s glands acts as an attrac-
tant to other turtles. Thus, the larger the aggregation becomes, the
stronger its attractiveness to other, still isolated turtles in the
vicinity, and eventually the entire population forms one huge ag
gregation. This may account for the fact that, from available data,
the arribadas in the old days, which numbered up to about 40,000
animals, were the only manifestations of nesting activity during
the entire season, there apparently being no isolated nestings on
other days. On the other hand, with today’s decimated population,
703
www.Ebook777.com
some small or medium-sized aggregations may form, but none be
comes large enough to attract all of the individual turtles in the
area; consequently nesting has become more diffuse, and
although small arribadas still take place, there is also a significant
amount of nesting by isolated individuals or small groups.
Nesting of most species of sea turtle has been shown to follow a
rather rigid schedule controlled by the internal cycles of the ani
mal; thus green turtles nest about every fourteen days, while
leatherbacks nest about every ten days during the season, and if
the turtle is frightened back to the sea before it can nest, it will re
appear rather promptly to try again—either later the same night,
or on the following night, in most cases. By contrast, ridleys have a
very flexible cycle, if indeed they have one at all, and it seems that
the animals can hold their fully-shelled eggs within their oviducts
until weather conditions are suitable for nesting. Optimal weather
conditions may be defined as cloudy, relatively cool days with a
strong north wind blowing, and these are sufficiently infrequent
that the turtles may have to wait several days or even weeks
before they can nest.
Relatively few data are available on the number of times a rid-
ley may nest during a season, but it appears that the maximum is
three, and it is probably more common for an individual turtle to
nest only once or twice. On the other hand, at least a proportion of
the nesting population will breed in successive years; this is ex
ceedingly rare for sea turtles of other genera.
A nesting ridley contrasts sharply with a nesting green turtle;
whereas the green moves ponderously, takes frequent rests, and
may be out of the sea for as much as two or three hours, the ridley
will trundle out of the sea with a great display of animation and
will march determinedly up to the dune area without a halt or with
only brief moments of rest. Having reached the dry sand above the
high tide mark, the turtle usually thrusts Its snout Into the sand
and, with powerful movements of its neck, appears to make some
kind of sensory appraisal of the sand. This is usually called ‘sand
smelling, ’ and indeed the action may well be olfactory in nature;
but it is also possible that the turtle is determining the texture or
water content of the sand.
The body pit, excavated so laboriously by the green turtle,
occupies the nesting ridley for only a few moments; after a few
simultaneous sand-sweeping movements of the fore flippers, these
flippers become firmly anchored in the sand and do not move
again until the eggs have been laid and covered up. The nest cavity
is, as usual, excavated entirely with the hind flippers. The turtle,'
working almost without interruption, inserts the flippers alter
nately into the ever-growing cavity, lifts out a ‘handful’ of sand,
and lays it down, while the hind part of the shell is moved over to
704
bring the opposite flipper into place. The sand removed by a par
ticular flipper action is not flicked aside (or forward) until just be
fore that flipper is about to be re-inserted into the cavity for the
next excavation. During the excavation the nuchal tendons appear
tense and prominent. The eyes are kept open; sand adheres to the
moist area around the eye, but the ‘tears’ are not nearly as copious
as those of nesting green turtles or leatherbacks.
When the nest cavity is completed, the flippers are splayed out
wide apart and oviposition commences. The cavity is now com
pletely concealed by the rear overhang of the carapace, though it
is possible to remove sand from the hind wall of the cavity if one
wishes to see the eggs dropping. At this stage the eyes are partially
or completely closed; there is considerable buccal action and
some partial opening of the mouth, and the deep breathing is clear
ly audible. As each batch of one, two, or three eggs is dropped, the
head and shoulders are withdrawn slightly. The actual dropping of
the eggs, which number between 50 and 160 (average 110), takes
only ten or twelve minutes. The eggs are white and spherical,
measure from 1.6” to 1.8” in diameter, and weigh, on average,
about one ounce each.
When the eggs have all been laid, sand is pulled into the cavity
with alternating curling-round strokes of each hind flipper. When
the eggs have been covered, the turtle commences a very charac
teristic ‘sand-thumping’ procedure that is peculiar to the genus
Lepidochelys. With eyes open and the distal parts of the foreflip
pers firmly anchored in the sand, the turtle rocks its shell vigor
ously from side to side so that each lateral margin of the shell is al
ternately brought into violent contact with the sand. The resulting
rapid thumping noise is audible at some distance. In between
bursts of four or five thumps the turtle pulls sand toward the tail
with the hind flippers. Eventually the fore flippers too throw sand
backwards between bursts of shell rocking. Finally, the shell-rock
ing is replaced by a curious series of movements in which sand is
thrown backward with simultaneous movements of one front flip
per and the opposite hind flipper. Several movements with such a
combination of flippers take place, then the opposing pair come
into action. Such a series of movements is occasionally interrupted
by simultaneous sand-flinging movements of the two front
flippers. These movements carry the turtle away from the nest
site along a sinuous path. Eventually the movement evolves into a
simple walk which takes the turtle back to the sea. The entire
process, from leaving the sea to re-entering it, does not usually
take more than 45 or 50 minutes, and even during the early stages
of the process the turtle shows little response to human observers
even at close quarters.
The incubation period for the eggs of this species under
natural conditions ranges from 50 to 70 days. Most emergences
705
www.Ebook777.com
take place between 53 and 56 days after laying. Chavez found the
fertility percentage for eggs transferred to a hatchery in 1966 to be
65.4, the eighteen clutches observed showing individual percen
tage fertility ranging from 43.0 to 95.5. Emergence usually takes
place in the early morning hours, during or shortly after dawn.
Apart from the now well-known mass nesting site near Rancho
Nuevo, Tamaulipas, very small numbers of nesting ridleys have
been reported on Padre Island, Texas, and one or two points on the
coasts of Veracruz and Tabasco. However, perhaps the most
bizarre re-nesting recovery of any sea turtle to date was that of a
ridley which was tagged while nesting at Rancho Nuevo in 1966,
and found re-nesting in July 1971 at Guachaca Beach, ten kilo
meters west of Buritaca on the Santa Marta Peninsula of Colom
bia! This area is hundreds of miles beyond the thitherto recorded
furthest wanderings of the species, and the fact that the animal
was nesting is even more remarkable.
Tagging experiments both by Humberto Chavez and by myself
have demonstrated that ridleys, after nesting in Tamaulipas, may
swim either north or south along the shores of the Gulf of Mexico,
there being a concentration of recoveries from the vicinity of
Laguna del Carmen, Campeche and also from the northern part of
the Gulf, especially near the mouth of the Mississippi River. Apart
from the above-mentioned nesting in Colombia, mature examples
of this species have never been found outside the Gulf of Mexico.
Nevertheless, immature specimens are most commonly found
along the Atlantic seaboard of the United States, and small ridleys
have been found washed up on the coasts of the British Isles on at
least twenty-five occasions.
It seems possible that newly-hatched ridleys embark on a
several-year-long trip that takes them well beyond the limits of the
Gulf but which eventually returns them to the nesting beach. Since
young ridleys are found in considerable numbers even as far north
as New England and the animals there appear to be in healthy con
dition, it is likely that these northern records do not necessarily
represent individuals permanently lost to the breeding population,
as has often been supposed in the past. As well as we can re-con-
struct from the observed distribution and size range of immature
ridleys, it appears most likely that after a ridley hatches and
enters the sea in southern Tamaulipas, Mexico, it swims actively
for some hours or days to eliminate the danger of being thrown
back on the shore, and then drifts more or less passively (perhaps
at times in association with floating mats of Sargassum weed) in a
clockwise direction in the Gulf of Mexico, passing with the Gulf
Stream south of Florida and up the eastern coast of the United
States. During this time it is feeding and growing, and by the time
it reaches the area offshore from New England it is large and
strong enough to become an active swimmer rather than a passive
706
drifter—by now it has a carapace length of about ten inches to one

foot. At this stage it reverses its previous direction of travel and
gradually migrates southward, so that by the time it is nearly
mature it has reached the Gulf coast of Florida, and a little later,
when maturity is reached, it has reached the food-rich areas off
the mouth of the Mississippi and near Laguna del Carmen, Cam
peche.
Some young turtles are apparently caught in the Gulf Stream
when they have reached the waters of the eastern United States
and are involuntarily carried across the Atlantic to the coasts of
Europe. It is likely that the size of such turtles when they arrive in
Europe is little greater than that when they left the Americas,
since the open sea is an alien and relatively non-productive habitat
which would offer them little chance of following a normal dietary
regimen. Thus, the very small ridley (99.7 mm in carapace length)
caught at Corvo, Azores, several decades ago probably left Ameri
can shores shortly after passing through the Florida Straits. These
turtles which stranded further north, which are almost always in
the 200 to 300 mm (8 to 12 inch) size range, had probably spent
several months moving up the Atlantic coast of the United States
before being caught by the Gulf Stream and transported to corres
pondingly more northerly points on the eastern shores of the At
lantic.
The diet of Kemp’s ridley is largely carnivorous, being known
to include crustaceans, jellyfish, molluscs (including cephalo-
pods), and fish. Gastropods and echinoderms are also eaten, but
the most plentiful individual items in the diet are probably crabs of
various species, including some very hard-shelled species. Kemp’s
ridley has very strong jaws and massive head musculature and
appears to have little difficulty in cracking even very thick clams
and crabs.
Ridleys do not commonly carry large numbers of barnacles,
as do many loggerheads and hawksbills, but a moderate propor
tion of adult females seen on the nesting beach show a few bar
nacles on the carapace and sometimes one or two on the head.
Mutilated turtles of this species are quite often found; typically,
one sees massive bites out of the rear part of the carapace, and
sometimes one or both of the hind flippers is also missing. This is
presumably the result of attack by sharks.
The ridley is reported to be of an hysterical and violent disposi
tion and to thrash and snap so implacably when caught and placed
on its back that it will expire from exhaustion rather than resign
itself to captivity. My only experience with this species has been on
the nesting beach; under these circumstances the turtles only rare
ly attempt to bite and may be turned on their backs for moderate
periods of time without fatal results.
The olive ridley, Lepidochelys olivacea, is of about the same
size as the Gulf species, but the two differ in a number of respects.
707
www.Ebook777.com
Essentially, olivacea is a more lightly built animal than kempi,
with a thinner and somewhat narrower shell and a small, more
lightly built skull whose individual bones do not become complete
ly fused together until comparatively late in life. The carapace of
olivacea is somewhat more elevated than that of kempi, with flat,
sloping sides and a flattened dorsal region. The costal scutes show
a remarkable variability; occasional specimens are found (espe
cially in the East Pacific) which show the same 5-5 costal count as
kempi, but typically the rearmost costals are split, giving a total
count of between six and nine (or occasionally more) costals on
each side. The central scutes too show some division, occasionally
numbering as many as eight or nine. It is quite common to find
olive ridleys which have different numbers of costal scutes on each
side, though costal counts differing by more than two are rare. The
most frequent number of costals in the Pacific and Indian Oceans
appears to be six on each side, while in the Guianas more individ
uals have seven than six. Kempi and olivacea are not distinguished
by their numbers of marginal scutes, but the mid-marginals are
much wider in kempi than in olivacea. The plastral scutes of oliva
cea are very stable, with the exception of the intergular which may
be absent, single, or double.
The jaws of olivacea are less massive than those of kempi and
lack the bony alveolar ridges of the latter species. The width
across the narrowest part of the pterygoid bones is greater, which
would indicate that the temporalis muscles which close the jaws
are smaller. The bony process on the lower jaw to which these
muscles attach is also smaller in olivacea than in kempi. The
orbits of olivacea are distinctly larger than those of kempi, pre
sumably because olivacea normally nests by night. In color there
is little difference between the two species; the oft-repeated dis
tinction which alludes to the grey color of kempi and the olive-
green color of olivacea is a fallacy which stems from a comparison
of juvenile kempi with adult olivacea. However, mature olivacea
do have a tendency to be a little darker than mature kempi.
The olive ridley is an exceedingly widespread species, being
found in the tropical waters of the Pacific, Indian, and South
Atlantic Oceans. In the East Pacific, the nesting range of the
species extends from the vicinity of Topalobampo, Sonora, to
Panama and possibly Colombia. The species is quite numerous in
Ecuador, but it has not yet been confirmed as a nesting species in
that country. I have seen numerous dead ridleys on the beach near
Salinas, Ecuador. Various sites of intensive nesting are known;
these are discussed below. The olive ridley is not known from the
mid-Pacific islands, though it has been recorded in the Yap and
Marianas Districts of Micronesia, and is widespread in the West
Pacific area; the species breeds in northern Australia, between
Gove Peninsula and Coburg Peninsula, and on Bathurst and Mel-
708
Lepidochelys olivacea,
the olive ridley. Seven
costals are visible;
this specimen came
from Guyana.
ville Islands, in many parts of Indonesia, in Malaysia (including

Sarawak), the Bonin Islands, southern Japan, and South Viet
Nam. In the Indian Ocean it is known from Karachi, the Sey
chelles, and Sri Lanka, but appears, to be very rare or absent from
most of the oceanic islands and from the Arabian Peninsula. In
East Africa, George Hughes has recently reported a substantial
population of this species in northern Mocambique and also in
northern Madagascar. In the East Atlantic, olive ridleys are
known to nest at least from Senegal to Zaire, while in South
America the species nests in considerable numbers in Surinam
and in smaller numbers in Guyana and French Guiana. Occasional
ridleys nest in Trinidad and possibly in parts of Venezuela and
Brazil.
The nesting process of olivacea shows no substantial differ
ence from that of kempi; this species also shows great urgency
during its ventures on to land and when nesting is not usually out of
the sea for more than about 45 minutes. As with kempi, the olive
ridley is relatively imperturbable when on land, and it is often pos
sible to tag the turtles as they leave the sea without frightening
them back. Indeed, on the Pacific coast of Honduras, I was
amazed to see the manner in which the local people obtained the
eggs of this species. Every night there were more ‘hueveros,’ or
709
www.Ebook777.com
egg collectors, than turtles on the beach, and when a turtle
appeared at the edge of the sea, several people would make a rush
to pick it up. The winner would carry the turtle to a pit in the sand
that he had prepared beforehand, and he would drop the turtle
quite heavily into this pit. The turtle would lie immobile for a
while, and sometimes would make for the sea immediately when it
came to its senses. But other times it would start the stereotyped
nest digging action with the hind flippers, which would be followed
by oviposition. As soon as all the eggs had been laid, the turtle
would be pushed aside, where it would go through useless cover
ing-up motions while its eggs were collected. Uncooperative tur
tles which tried to return to the sea after being dumped in the pit
were simply brought back time and time again until they started
digging an egg cavity. I do not believe any other species of turtle
would submit to this interruption of its normal nesting prelimin
aries, though I did once turn over a leatherback before it had a
chance to nest, and when I turned it back by daylight several hours
later it started digging a nest cavity.
It is noteworthy that olivacea, like kempi, shows a strong pref
erence for nesting when strong winds are blowing off the sea; but
in contrast to kempi, the nesting of olivacea usually takes place by
night.
Until the late 1960’s, it was believed that olivacea showed a
complete contrast to kempi in emerging singly to nest on any suit
able beach, rather than forming massive nesting aggregations.
However, it is now known that olivacea forms arribadas at several
remote places on both sides of the American continent, and that
some of these arribadas are even bigger than were those of kempi
thirty years ago, before the latter species was decimated. In the
Atlantic, only one olive ridley arribada site is known; this is the
small beach called Eilanti, on the western part of the coast of Suri
nam, near the mouth of the Marowijne River which separates Suri
nam from French Guiana. The arribadas here are not particularly
large—up to about 500 turtles on a peak night—but the beach is
very small (about half a mile long), so the concentration is quite
impressive. Moreover the manifestation always occurs at the
same place, and at approximately predictable times (at fortnight
ly intervals in June and July), so it is much easier to meet up with
this arribada than with any other. For many years the local Carib
Indians collected virtually every egg laid on this beach, and the
turtle population was rapidly headed for extinction. In the 1967 and
1968 seasons, with the help of financial support from the World
Wildlife Fund, I was able to purchase the entire seasons’ supply of
eggs from the Indians and to hatch them out—in a hatchery the
first year, in situ the next year—and thus, for the first time for
many years, substantial numbers of hatchlings reached the sea
safely. The Surinam Government then took the initiative and in
cluded the beach at Eilanti in their National Park system, though
under a special arrangement some egg collection was still permit
ted. However, since the numbers of ridleys at Eilanti showed a
marked decline in 1970 and 1971, collection of ridley eggs in Suri
nam is now completely prohibited.
As mentioned earlier, the arribadas at Eilanti follow a some
what predictable cycle; there is a ‘background’ level of between
about 2 and 20 ridleys nesting each night throughout the season,
but, usually in early June, the numbers will build up in the course
of two or three nights to a peak which may number as many as 500
turtles. During the following two or three nights the numbers will
decline again to a background level. The process is repeated at
least three times during the season, with intervals of about two
weeks between peak nights. However, if there should be no wind
during an anticipated peak night, no turtles, or very few, will nest.
The big arribadas show a strong tendency to appear in the first
half of the night, on nights when the tide is rising at dusk and fall
ing at midnight. This is perhaps a mechanism to allow those tur
tles which may be accidentally stranded on the enormous mud
bank in front of the beach to be re-floated by the incoming dawn
tide without having to endure a period of exposure to the tropical
sun.
In Mexico, ridleys may form enormous arribadas, numbering
many thousands of animals, at several points on the Pacific coast.
These points include Playon de Mismaloya in Jalisco; between the
village of San Luis la Loma and the rock known as La Piedra de
Tlacoyunque in Guerrero; at Playa Escobilla between the coastal
towns of Puerto Angel and Puerto Escondido in Oaxaca; and near
the Rio Apiza mouth on the Colima/Michoacan State Line. All of
these arribadas are impressive spectacles and may last for two or
three days and nights. The largest arribada on definite record was
seen at Escobilla, Oaxaca, between August 7 and 10, 1968; it is esti
mated that a total of 80,000 turtles nested during this period. How
ever, because these arribadas do not always take place in exactly
the same part of the beach, and because they are highly unpredic
table as to time, they are not easy to meet up with. Nevertheless
local people are able to sense when the turtles are ready to nest,
and they move down to the beaches and indulge in orgies of egg
collecting. Fortunately the Mexican turtle conservation service
maintains parties of biologists, fisheries inspectors, and fully
armed marines on these beaches during the nesting season, and
thus the worst of the destruction is avoided.
It has recently come to light that arribadas of enormous size
also occur in the remote northwestern part of Costa Rica near the
Nicaraguan border. Here, on Playa Nancite and Playa Ostional,
711
www.Ebook777.com
arribadas fully as large as any in Mexico come ashore on unpre
dictable dates in the latter part of the year. The arribadas them
selves are difficult to meet up with, but from a low-flying aircraft
one can see the enormous numbers of turtles aggregating at sea in
preparation for their massive synchronized nesting effort. Photos
of the daylight arribada on Nancite Beach in late 1971, taken by
David Hughes of the University of Miami and published in the Oct
ober 1973 National Geographic and the September 1973 Audubon,
must rank among the most spectacular turtle pictures ever taken.
A brief notice in the May, 1976IUCN Bulletin suggests that one
of the largest arribada sites of all, and the only one known in the
Indian Ocean, may have escaped discovery until now. This
rookery is located on the coast of Orissa, India, and according to
the notice, Dr. H.R. Bustard marked a total of 158,161 nesting
female olive ridleys during a three-week period in early 1976. Dr.
Bustard reported that until this year, between 1.5 and 2 million
eggs were taken every year, but that on his advice the government
of Orissa has completely banned the taking of eggs and has
revoked egg licenses.
The diet of the olive ridley is known to include small shrimp,
jellyfish, crabs, snails, and fish. In keeping with the lighter con
struction of the jaws of this species, the diet does not appear to in
clude as many hard-shelled organisms as does that of kempi. In
Ceylon, Deraniyagala often found the stomachs of olive ridleys to
be filled with vegetable matter (algae).
Clutch size of the olive ridley is similar to that of Kemp’s rid
ley; 928 clutches from Eilanti, Surinam, contained between 30 and
168 eggs, the average being 116.07. Sixty-five clutches from Pacific
Honduras ranged from 48 to 151, with an average of 106.00. The dif
ference in average values for the two areas may not be significant,
since the Honduras clutches were produced toward the end of the
nesting season and thus would include a higher proportion of
small, end-of-season clutches.
The migrations of olivacea throughout most of its range are
poorly known; however, stray individuals have been found in the
East Pacific as far north as Oregon and as far south as Chile, even
though the nesting range reaches north only to the central Gulf of
California and south to Panama. In the Guianas over 2000 ridleys
have now been tagged, in latter years with the help of the Surinam
Forest Service and the Dutch Army; nearly all of these turtles
were females nesting at Eilanti, Surinam. About fifty of these rid
leys have since been caught at sea. The highest densities of recap
tures are along the coasts of Surinam and French Guiana, but
there are a few records for Guyana and the north coast of Brazil,
and a definite concentration of recoveries around Trinidad and the
vicinity of Carupano, Venezuela.
712
Olive ridleys of between hatchling and mature size are almost

never caught; I have spoken with proprietors of turtle coopera
tives in Mexico who handle tens of thousands of olive ridleys
annually, yet they have never seen half-grown individuals. I can
offer no explanation for this phenomenon.
GENUS ERETMOCHELYS
The relationships of the hawksbill turtle, Eretmochelys imbri-
cata, are somewhat uncertain. The loggerhead and the two ridley
species form a natural grouping that is sometimes called the
Carettini, and the green turtle and the flatback are closely related
and are often placed in the subfamily Chelonini. But whether the
hawksbill falls more naturally into the Carettini or the Chelonini
has been the subject of rancorous discord. Archie Carr (1942, Proc.
New England Zool. Club, XXI: 1-16) listed the following points
which suggest that the green turtle should be considered apart
from the others, and that the hawksbill should be placed with the
loggerhead and the ridleys. Only the green turtle has: i) ptery
goids that are not deeply concave; ii) a short mandibular symphy
sis; iii) no anteriorly projecting boss for attachment of the pos
terior ectopterygoideus muscle at the symphysis of the quadrate
and prootic; iv) inner surface of the upper beak strongly ribbed
vertically; v) edge of the lower beak deeply dentate; vi) median
alveolar ‘tooth’ of the upper jaw connected with terminal ‘tooth’ by
sharp ridge; vii) numerous choanal papillae (only one, or at the
most two, in the other sea turtles); viii) only one pair of prefrontal
scutes (two pairs in the other species); ix) no lateral carapace
ridges in the hatchling; x) plastron of the hatchling with no dark
pigment (though dark pigment appears very soon after hatching in
the East Pacific green turtle); xi) only one claw on each fore flip
per; xii) three, rather than two, sacral ribs; xiii) the last peripher
al bone not entered by a rib lying between rib numbers 8 and 9
(numbers 7 and 8 in the other species).
Other authors have considered many of the peculiar features
of the green turtle listed by Carr to be morphologically or function
ally correlated, and have stressed certain points in which the
hawksbill resembles the green turtle in placing Eretmochelys in
the Chelonini. For example, both the green turtle and the
hawksbill have only four pairs of costal scutes, in contrast to the
loggerhead and the two ridleys, which have at least five pairs. The
first two species also have elongate coracoids, while in the latter
three the coracoids are only about as long as the scapulae. The
green and the hawksbill have eleven pairs of peripheral bones,
while the loggerhead and ridleys have at least twelve pairs. One
final feature is that the nuchal scute of Caretta and Lepidochelys is
in contact with the first costal scutes, while in Chelonia and Fret-
713
www.Ebook777.com
mochelys the nuchal is separated from the first costals. However,
this difference is surely correlated with the difference in costal
scute counts between the two groups; in the loggerhead and rid-
leys, the insertion of a small extra costal at the beginning of the
series causes a narrowing of the first central, which thus allows
the wide nuchal to make contact laterally with each of the first cos
tals. On balance, it seems most reasonable to consider the genus
Chelonia on its own and to group Eretmochelys, Caretta, and Lepi-
dochelys together. A glance at the highly distinctive hatchling of
the green turtle re-inforces this opinion, as does the mode of pro
gression of each of the species on land. Only Chelonia moves by
means of symmetrical, humping-forward heaves; the others all
have a normal tetrapod walk, with alternating movements of op
posed flippers. This difference in mode of progression has been
correlated with the different weights of the animals, since the
green is the largest hard-shelled sea turtle, and the enormous
leatherback also shows a Chelonia-type gait; but the difference
holds up even in the small East Pacific green turtle and the moder
ate-sized Chelonia depressa, both of which are smaller than log
gerheads and similar in size to Atlantic hawksbills.
The hawksbill is a small or medium-sized sea turtle; mature
females in Costa Rica and Guyana are about 31” to 36” in carapace
length; a series of ten from Guyana weighed from 98 to 165 pounds.
In the Arabian Peninsula hawksbills do not reach this size; four
teen specimens measured by Harold Hirth at Jabal Aziz Island, in
the Gulf of Aden, measured 25” to 28.5” in carapace length and
weighed from 78 to 110 pounds. This also appears to be the normal
size of mature hawksbills in the East Pacific. The heaviest known
hawksbill was caught off Grand Cayman and weighed 280 pounds;
the longest on record was one I found in French Guiana with a 37”
carapace.
The hawksbill is known throughout Latin America as carey.
This is one of those curious vernacular names which is used for a
single turtle species in many different countries, transcending
barriers of culture and even language. Another example is mata-
mata for Chelus fimbriatus, and perhaps also cahuama (for sea
turtles—usually the loggerhead) and jicotea (widely used for
Pseudemys).
The carapace of the hawksbill is relatively narrow and is
strongly serrated behind in all except very old individuals. The
scutes of the carapace are thicker than in any other turtle and
cover a bony structure that is relatively light and in which costo-
marginal fontanelles usually persist throughout life. The carapace
scutes are strongly overlapping in most specimens, in contrast to
those of other sea turtles, in which the scutes always meet edge-to-
edge. However, the overlap is not evident in the hatchlings, and in
714
very old individuals the free posterior edges of the centrals and
costals wear away and the scutes assume a juxtaposed condition.
The coloration of the dorsal scutes of the hawksbill is most attrac
tive, with radiating streaks of brown and black in an amber sub
strate. However, the pattern only assumes its full beauty in the
separated, polished scutes (“tortoise-shell”)—in living hawksbills
the carapace is often encrusted with barnacles and other epizootic
organisms. The plastral scutes of the hawksbill are clear yellow or
orange-yellow; they show a distinct overlap in younger specimens.
Four pairs of inframarginal scutes are present.
The head of the hawksbill is relatively very narrow, parallel
sided, and with the snout tapering to a point. The jaws are ridged
but are not hooked, and the term ‘hawksbill’ is thus somewhat of a
misnomer, though the bill is certainly bird-like. There are two
pairs of prefrontal scales. The scales of the head and forelimbs are
very distinct, each being very dark brown or black with a light bor
der. The soft skin of the ventral parts of the mature animal has an
orange tinge. The skin on the underside of the flippers of Atlantic
hawksbills is almost lacking in dark pigment, except toward the
outer ends.
The subspecies of Eretmochelys imbricata are still imper
fectly characterized. At the present time it is customary to call
Atlantic hawksbills Eretmochelys imbricata imbricata (type
locality: Bermuda) and to call Indian Ocean and Pacific hawks
bills either Eretmochelys imbricata squamata (type locality:
Singapore) or E.i. bissa (type locality: Red Sea). I suspect that at
least three subspecies of the hawksbill could be demonstrated to
exist: one in the Atlantic Ocean, one in the Indian Ocean and wes
tern Pacific, and one (which would need a new name) in the East
Pacific. The Atlantic populations seem to be characterized by a
relatively narrow carapace, large adult size, a dorsal scute pat
tern consisting of a relatively simple arrangement of brown and
amber, the scales on top of the head and on the upper surfaces of
the forelimbs dark brown in color with rather broad yellow mar
gins, and the upper mandible predominantly yellow, with relative
ly little dark pigmentation. The Indian Ocean and West Pacific
populations seem to be rather variable but tend toward a broad
carapace, rather small adult size, and a highly ornate pattern on
the carapace scutes, with several series of short radiating streaks
on each of the costals and centrals. The hawksbills of the East
Pacific are relatively broad, of small adult size, and have
intensely black scales on the upper surfaces of the head and fore
limbs, each'scale having only a narrow border of a pale yellow or
white color. The carapace scutes are ornate but heavily pigmented
with black, and there are also frequently black blotches on some of
the plastral scutes. Even within the East Pacific area, however,
715
www.Ebook777.com
there appears to be variation; for example, the top of the head is
usually uninterrupted black in specimens from Honduras, while
all the scales have light borders In the Galapagos Islands. Atlantic
hawksbills seem to be characterized by particularly jagged serra
tions on the hind margin of the carapace and may also show a les
ser development of the dorsal keels than do Indo-Pacific speci
mens.
The hawksbill is a confirmedly tropical animal. There are only
two records of the species nesting in the mainland United States
(on Juno Beach, Florida), and only one stray specimen is definite
ly known to have reached European Atlantic waters. The pre
ferred, though not exclusive, habitat of the Atlantic hawksbill con
sists of coral reefs and relatively shallow, rock-bottomed coastal
waters. Hawksbills are also found in mangrove-bordered shallow
inlets. The nesting beaches are located on remote oceanic islands,
on offshore cays, and on mainland shores. In most areas the
hawksbill is what may be called an opportunistic nester, emerging
to lay at any time during a long season on any beach convenient to
its feeding grounds. Long migrations have not been recorded for
this species, but two female hawksbills tagged at Tortuguero,
Costa Rica, have been recovered respectively 285 and 325 miles to
the north. The hawksbill is the most abundant sea turtle on most of
the Lesser and Greater Antilles, but nevertheless the population
density in this area is rather low, nesting is sporadic, and no
rookeries are known.
A few hawksbills nest on the Caribbean coast of Costa Rica-
some among the green turtles at Tortuguero and possibly slightly
greater numbers further to the south. There appears to be a defi
nite rookery for the hawksbill at Chiriqui, on the western part of
the Caribbean coast of Panama. The species has apparently been
almost exterminated as a nesting animal in Colombia and prob
ably in most of Venezuela also. However, on Shell Beach, in the
Northwest District of Guyana, one or two hawksbills may be found
nesting on most nights in July and early August. A very few nest at
Bigisanti, Surinam, and each season, while tagging leatherbacks
in French Guiana, we find between one and five nesting hawks
bills. The habitat here is marginal, however, the sea being shallow
and muddy, with no rocks or coral. Young hawksbills are quite
often seen around Devil’s Island, where the water is clear and the
sea bottom rocky.
The hawksbill is apparently still reasonably plentiful near
some of the remote West Caribbean Islands such as Old Provi
dence and San Andres, and around the Serrano and Serranilla
Banks; however it is universally heavily exploited. Although vir
tually gone from the mainland beaches of Colombia, the species is
still found on some of the offshore cays, especially Cayo Roncador.
716
There are also some beaches and islands off the east coast of Nica
ragua where small numbers of hawksbills still nest. Perhaps the
most important one left is the mainland beach near Cocal on the
southern part of the east coast of Nicaragua, but other hawksbill
beaches in Nicaragua listed by Bernard Nietschmann (in manu
script) are: the Set Net Peninsula from Parakeet Point to about 15
kilometers south of Tasbapauni; the Set Net Cays (Water Cay,
Grape Cay, Wild Cane Cay, Crawl Cay, Baboon Cay, Lime Cay,
Vincent Cay, Black Mangrove Cay, and the two Tangwira Cays);
the Asking Cay area (Big Asking Cay and the southern Savanna
Cay); and the King’s Cay area (Swirri Cay and Big King’s Cay).
This may seem like a lot of hawksbills, but in fact the population
density is so low that the typical pattern of exploitation is for a
turtler to find a fresh nest on a beach by chance and to return to the
same beach 14 or 15 nights later in the hope of finding the same tur
tle re-nesting.
Hawksbills exist, but are scarce, in the Gulf of Mexico. They
are caught off the coasts of Campeche and Veracruz and in former
times nested on the island of Sacrificios, near the city of Veracruz.
The species is also found in Bermuda, the Bahamas, and Cuba, and
is occasionally caught in the Florida Keys.
In the eastern Atlantic the hawksbill is known from the Medi
terranean, though nesting has not been recorded. The species is
found in West Africa, but I know of no studies of the species in this
area. Museum specimens exist for Senegal, Gabon, Cameroon,
Sierra Leone, Ghana, Liberia, and the Cape Verde Islands. Nest
ing occurs on Rolas and Sao Thome Islands. In the Indian Ocean a
hawksbill rookery is known on the small island of Jabal Aziz, off
the south coast of the Arabian Peninsula near Aden. Harold Hirth
found seventeen nesting hawksbills on the one mile beach on this
island on February 1,1967, and sixteen on the night of February 8.
Peter Walczak and Wayne Kinunen have recently demonstra
ted that certain islands in the Persian Gulf attract large numbers
of nesting hawksbills. The best location found was a 600 meter
beach on the eastern shore of Shitvar Island, where over 250 nests
were found when the island was visited in June, 1971. Shitvar is un
inhabited and has no domestic mammals, so there is hope that the
turtle population can continue unmolested. Shitvar Island is only
three square kilometers in area and is located two kilometers from
the eastern tip of Shaikh Shu'aib, or Lavan Island. Lavan is also an
important hawksbill rookery; 218 nests were found there in June
1971, the majority being hawksbill nests and the remainder nests
of the green turtle. Hawksbill nesting also takes place on Qeshm
Island, in the Strait of Hormoz at the mouth of the Persian Gulf.
Despite the abundance of hawksbills in the Gulf of Aden and
the Persian Gulf, there is no record of the species breeding in Pak-
717
www.Ebook777.com
istan. The hawksbill used to be very abundant around Socotra, but
at least eighteen centuries of exploitation have understandably
made it scarce today in this region.
Hawksbills are found in the Seychelles Islands and are said to
be very common at Passe Houareau and Main Channel. A few are
caught at Aldabra Island. Jack Frazier examined two specimens
caught there during his seven-month stay on the island and found
that they agreed with Atlantic hawksbills in most characters
that have been used to distinguish the various races—i.e. there
was a median keel on the last four centrals; also lateral ridges con
verging posteriorly were only present on the last central; no
lateral keels; plastral keels not prominent; margin of carapace
serrate; carapace rather narrow, with parallel sides. However,
the turtles had the intense black pigmentation of the head and the
dorsal parts of the flippers, which is considered characteristic of
certain Pacific populations. Clearly the geographic variation of
the hawksbill requires much more study. To complicate matters
further, Hornell (1927, The turtle fisheries of the Seychelles
Islands. London, H.M.S.O.) reports that the Aldabra hawksbills
often have the highly prized ‘blonde’ carapace scutes, in which
each scute has extensive unpigmented areas. A color photograph
sent to me by George Hughes showing a hawksbill from Antonio
Enes, northern Mocambique, is also of the ‘blonde’ type. In Mo-
cambique, the hawksbill is reported to be as common as the green
turtle in the coral reef area north of Antonio Enes. Nesting is said
to take place abundantly on the islands of Quirimba, Singar, and
Mefunvo near Ibo, north of Porto Amelia, and a little nesting takes
place on the mainland of northern Mocambique. Nesting has not
been recorded on the mainland of Madagascar but is reported to
take place on some of the smaller islands off the west coast.
In the northern and eastern Indian Ocean and in the West
Pacific, the hawksbill probably nests in a great many localities,
but specific literature records are rather few. The species nests in
Ceylon and is said to abound near Celebes. I.S. Suwelo lists these
other Indonesian Islands from which the species is known: Suma
tra, Natuna, Borneo, Djawa, Madura, Sumba, Flores, Ambon,
Banda, Obi, Temato, Kei, Aru, and Irian (or New Guinea). G.S. de
Silva has recorded nesting hawksbills in the first three months of
the year both on the mainland of Sabah as well as on the islands of
Pulau Selingaan, Pulau Gulisaan, Pulau Bakungaan Kechil, Pulau
Bilean, Pulau Tegapil, Kota Belud, Pulau Mantanani, and Pulau
Lankayan. Robert Bustard has recently announced the discovery
of what must be the most important hawksbill nesting beach in the
world. This is situated on Long Island (approximately 10°S, 143°E)
in the Torres Strait, off northern Australia. The turtles here nest
both by day and night, and 76 nested within one 24-hour period in
718
July 1972. Despite severe cyclone weather which completely pre

vented any nesting on certain nights, no fewer than 210 hawksbills
nested during a nine-night period.
Among the innumerable islands of the tropical Pacific, hawks
bills appear to be widely distributed and to nest in many places,
though up to now rookeries have not been recorded. The species is
fairly common in the Ryukyu Islands. Hawksbills are known to
nest regularly on a small island off the coast of Western Samoa,
and those eggs which escape egg-stealers are being hatched and
the young turtles released by Peace Corps volunteers. Sporadic
hawksbill nesting is known on several of the Fiji Islands, including
Laucala, Matagi, and Nanuky Lailai. In the islands of the U.S.
Trust Territory (Micronesia), hawksbills are widely distributed
but generally rather rare except in the Palau District. In the lime
stone islands area of the Palau lagoon, however, hawksbills are
still common and nest during the summer months on many islands
including Aulong, Ngeangas, Ngobadangel, Unkaseri, Abappao-
mogan, Eomogan, and Ngerugelbtang.
Hawksbills exist, but are very rare, in the Galapagos Islands.
There is also one nesting record for Halawa Beach, Molokai,
Hawaii. On the mainland shores of the East Pacific the hawksbill
is known to nest diffusely along the coast of Mexico, from Nayarit
to Chiapas. There is a zone of moderately concentrated nesting
near the southern tip of Baja California, and nesting also takes
place in the Tres Marias Islands, including Isla San Juanito. Nest
ing is also known on islands in the Gulf of Fonseca, Honduras, and
on the coasts of Nicaragua and El Salvador.
Hawksbill nesting almost always takes place by night, but has
not yet been shown to have lunar or tidal correlations. The animal
walks with a typical tetrapod gait, and with its relatively long,
mobile, and partially retractile neck has much less of the auto-
maton-like aspect of a beached green turtle. The female hawksbill
is highly circumspect as she comes ashore and makes both a vis
ual appraisal of the beach with much craning of the neck and look
ing around, as well as an olfactory appraisal of the substrate by
thrusting her snout strongly into the sand in front of her. The turtle
may walk about extensively before selecting a nest site (several
hundred yards in some cases), but once the site is chosen, the
‘body pit’ phase of excavation is symbolically completed with just
a few strokes of the fore flippers before the serious business of ex
cavating the egg cavity begins. The alternating digging strokes of
the hind flippers, with each foot insertion preceded by a strong for
ward ‘flick,’ proceeds exactly as in other sea turtles. However,
during actual oviposition the hind flippers are splayed out widely
rather than being employed to cover the egg cavity as in the green
turtle and the leatherback, and each egg deposition is accom-
719
www.Ebook777.com
panied by an upward curling of the hind edges of the hind flippers.
The covering up of the eggs and subsequent disguising of the nest
site show no great peculiarities. The return to the sea is compara
tively rapid, though if the animal is very tired it may rest periodic
ally or make a few green-turtle-like breast-stroke movements of
the flippers.
The Atlantic hawksbill lays more eggs, on average, than any
other turtle. Fifty-seven nests examined by Archie Carr and his co
workers at Tortuguero, Costa Rica, contained on average 161.1
eggs, the overall range being 53 to 206. Seven nests that I counted
in Guyana contained from 139 to 176 eggs (mean 158). In the Mis-
kito Keys area of Nicaragua, local turtlemen report that the
hawksbill lays about three times during a season, at intervals of
about 15 days, and that the number of eggs in each clutch dimin
ishes steadily; the first nest may number 160 to 180 eggs, the
second about 130, and the third about 60. The largest hawksbill
clutch on record contained 221 eggs and was deposited by a French
Guiana turtle in 1972. Clutches are much smaller in the Indian
Ocean; Hirth found five nests at Jabal Aziz Island, in the Gulf of
Aden, to number 96, 114, 127, 99, and 103 eggs respectively. More
over each of these clutches contained a rather high proportion (19
to 39 percent) of undersized, yolkless eggs. Deraniyagala found 115
eggs in a Ceylon nest.* The eggs are about 1.6” in diameter and are
thus indistinguishable on the basis of size from ridley eggs. In
some areas the eggs have a slight pink tinge. The young emerge,
usually by night, after about two months. The hatchling is brown in
color, has three dorsal keels and two ventral keels, and lacks both
the imbricated scutes and elongated, narrow snout which are so
characteristic of the half grown and adult turtles. The mean cara
pace length at hatching is 1.67”, at least in the Caribbean.
It is not known with certainty whether hawksbills nest every
year or every second or third year. One turtle tagged while nesting
at Tortuguero was seen re-nesting three years later, and another
six years later. The hawksbill is a largely carnivorous species and
feeds upon the wide variety of organisms that its tropical reef
habitat presents. Stomach analyses have revealed that sponges
figure prominently in the diet, but coelenterates, ectoprocts, echi-
noderms, gastropod and bivalve molluscs, barnacles, crusta
ceans, ascidians, and fish have also been reported. The narrow
snout and head and elongate neck of the hawksbill presumably
represent an adaptation toward extracting food from rocky cre
vices, and the heavily comified shell helps protect the animal from
excessive buffeting against rocks and coral heads. Freshly caught
hawksbills are pugnacious and bite hard. However, the animal
when on the nesting beach does not attempt to bite even when
molested.
*On Cousin I. (Seychelles) a female laid 252 eggs.
720
John Steinbeck, in his book The Log of the Sea of Cortez,

records the capture of a hawksbill turtle off the southern coast of
western Baja California. The entire digestive tract of the animal
was crammed with small bright red rock-lobsters (Pleuroncod.es
planiceps). The active peristaltic effect of the posteriorly-directed
horny spikes which lined the digestive tract, whose movement was
still visible in the freshly killed animal, had the function not only of
macerating the food, but also of moving it down toward the stom
ach. This mechanism is probably also operative in other sea turtle
species.
The beautiful scutes of the hawksbill turtle have been prized
by man for so long that the species is rapidly disappearing from
many, if not most, of the places where it was formerly abundant.
The scutes are removed by application of heat to the live or dead
animal, and subsequent heat treatment allows the plates to be
shaped and even welded together. The product is known as tor
toiseshell and is used for making spectacle frames, brooches,
combs, and even small boxes. Fortunately it is now illegal to
import the product into the United States, and it is hoped that soon
the European countries will follow suit and ban the importation of
this purely luxury product which is derived from a highly threat
ened species. At the present time very large numbers of young
hawksbills are killed and stuffed and their entire shells polished up
to make curios which may be hung on the wall. Russell Mitter-
meier estimated that in Mexico as many as 2000 stuffed juvenile
hawksbills are sold annually in the city of Veracruz alone. A recent
report by I.S. Suwelo indicated that on the island of Tidung, in the
Seribu Archipelago in the Java Sea not far from Jakarta, most of
the i milies raise hawksbills from hatching to an age of up to about
two months, entirely for the curio trade. Each family keeps be
tween about ten and one hundred turtles, feeding them on rice and
kitchen scraps as well as chopped fish. In 1972 it was estimated
that there were about 4600 young hawksbills in captivity on the
island. The turtles are sometimes sold alive for the equivalent of
two to four dollars, but others are sold in a stuffed and formalin-
preserved form for about six dollars.
Hawksbills are also raised for souvenir purposes by aboriginal
people in northern Australia. Many of the inhabited islands of the
Torres Strait have small turtle-raising projects, which it is hoped
will eventually give them a steady source of outside income. These
projects were visited recently by Dr. Archie Carr, who was aston
ished to find that the hawksbills from different islands in the Tor
res Strait frequently had quite distinct shell patterns almost from
hatching. Since in many cases turtles showing such differences
had been raised on the same farm under identical conditions, the
conclusion is inescapable that the situation represents a spectacu-
721
www.Ebook777.com
lar example of micro-evolution between adjacent populations iso
lated solely by the strength and single-mindedness of the urge of
the female—and male—turtles to return to their natal beach for
breeding. The various populations appear to have similar shell
shapes, but the pigmentation of the scutes of the carapace is some
times coarsely variegated, sometimes finely variegated, and
sometimes relatively plain, each scute simply having a relatively
dark central or posterior area, with the areas of relatively recent
growth being lighter. The Australian turtle ranchers get a much
better price for their stuffed hawksbills than do the inhabitants of
Tidung Island.
The flesh of the hawksbill is little favored for human consump
tion in most areas and indeed is occasionally fatally poisonous;
Deraniyagala recorded a case of twelve people in the village of
Karuppankudiyiruppu in Sri Lanka dying from eating hawksbill in
1888. However, hawksbills are eaten in Guyana and Campeche, al
though in neither locality is the shell utilized; and in some few
places, such as Cayman Brae and Old Providence Island, the
hawksbill is actually considered more tasty than the green turtle.
GENUS DERMOCHELYS
The leatherback turtle, Dermochelys coriacea, is the sole spe
cies included in the family Dermochelyidae. Indeed, so divergent
is this species in so many aspects of its anatomy that it is often
placed in a separate suborder, the Athecae, to contrast it with all
other turtles that are placed in the suborder Thecophora. However,
the leatherback does agree with the other sea turtles in certain
aspects—such as the roofed-over skull and many aspects of its
nesting behavior—and the family Dermochelyidae is thus con
sidered related to the Cheloniidae rather than representing a com
pletely separate adaptation of turtles toward a marine environ
ment.
The leatherback is easily the largest species of turtle in the
world; nevertheless its maximum size has been exaggerated. The
literature abounds with unconfirmed records of eight or even nine
foot long leatherbacks, but most museum specimens of adult
leatherbacks have carapace lengths of only about five feet or a lit
tle more (approximately another sixteen inches may be added to
give a figure for the total length, or a few more inches still for the
total length of the long-tailed male). I have measured more than
2500 mature female leatherbacks on the nesting beach in French
Guiana and have found some bigger than any museum specimens I
have seen, but I have yet to encounter a six foot leatherback, al
though the three biggest each had shell lengths of exactly 71
inches. The median carapace length, at least in the West Atlantic,
is about 61 inches. There are few reliable records of the weights of
722
adult leatherbacks, but Agassiz’ statement that he had seen speci

mens weighing over a ton is usually assumed to be an exaggera
tion. Having established the weight of a rather small (58”) leather
back as 651 lbs., I would suggest that the average is about 800 lbs.
and the maximum about 1300 lbs.
The leatherback is immediately distinguishable by the
absence of all cornified epidermal structures; it has no scutes on
carapace or plastron, no scales on the skin, no claws, and not even
flattened homy plates in the jaws. The carapace has the texture of
hard vulcanized rubber and is somewhat flexible to the touch; the
plastron is very soft. However, the carapace gains a considerable
amount of rigidity by being raised into a number of longitudinal
ridges. The median ridge, particularly toward the rear, is the high
est, but there are two others on each side between the median one
and the lateral margins of the shell. The animal has a rather
barrel-shaped form with no sharp angle between carapace and
723
www.Ebook777.com
plastron. There are feeble suggestions of five longitudinal ridges
along the plastron.
Anteriorly, the shell of the leatherback grades almost imper
ceptibly into the massive neck and shoulder region. The head is
large, with eyes that open in a strongly oblique, almost vertical
plane. The lower jaw has a median hook and the upper jaw has a
pair of strong cusps flanked by deep notches. The jaws are sharp-
edged but not particularly strong. The front flippers are extremely
long—the stretched-out flippers of an adult specimen may span as
much as nine feet. The track of a leatherback on a beach usually
measures six to seven feet in width. Posteriorly, the carapace has
a moderate overhang and tapers strongly to a blunt point. The hind
flippers are broad and are connected to the tail by folds of skin.
In color the leatherback is predominantly black. However,
most individuals have at least some white spots on the shell and
skin, and frequently the white spotting is very dense. White, or
rather pale pink, predominates on the ventral surface of the ani
mal. The spots sometimes show a pink or bluish tinge, but are
never yellow, as has sometimes been reported. All of the mature
Leatherback turtles on a nesting beach in Trengganu, Malaysia. Photo by

Malay Information Center.
724
female leatherbacks that I have seen had a pink area, of very vari
able extent, on the crown of the head.
Skeletally the leatherback is most peculiar. In the carapace,
the butterfly-shaped nuchal bone is well developed, but neural,
pleural, and peripheral bones have completely disappeared. The
ribs are well-defined but are thin and contribute little to the
mechanical strength of the carapace; they are unique among tur
tles in lacking lateral coalescing flanges and in being merely in
loose juxtaposition with the carapace, rather than embedded in it.
Epi-, hyo-, hypo-, and xiphiplastra are present but are reduced to a
thin rim of bones around the edge of the plastron. The ‘shell’ of the
leatherback is about 1.5” thick and is composed largely of tough,
greasy cartilage. Just under the surface of the carapace, however,
there is a layer composed of thousands of small, polygonal bones
which form a continuous mosaic. The longitudinal ridges of the
carapace are composed of somewhat enlarged, tectiform, inter
mittently tuberculate mosaic bones. The continuity of the mosaic
breaks down in the region of the bridge, and in the plastron only a
few isolated polygonal bones are present, concentrated along the
ridges. These mosaic bones are probably neomorphs, though some
consider them homologous to the epidermal bones present in the
backs of crocodiles or the front legs of tortoises.
The most massive bones of the leatherback body are the scap
ulae and the rather short but very thick humeri; it is to these bones
that the enormous swimming muscles attach. The skull is sturdily
constructed dorsally, but the palatal area is quite flimsy. The pari
etal bones lack the descending processes present in cheloniid sea
turtles. There is no secondary palate, and palatine fenestra also
are absent. The quadrate does not completely enclose the stapes,
being open posteriorly. The vomer meets the premaxilla, separa
ting the palatines. The basisphenoid does not touch the palatines,
and the maxilla is separated from both the pterygoid and the quad-
ratojugal on each side. The orbits are very large, and the skull roof
extends even further back than in other sea turtles.
A peculiar feature of the leatherback skeleton is its retention
of essentially embryonic characteristics into adult life. In even a
big, mature leatherback, the limb bones retain extensive cartilagi
nous ends, while the pelvis contains so much cartilage that the six
separate bones of a dry preparation do not seem to fit together at
all. The skull also falls apart completely on maceration. Moreover
the entire skeleton, as well as the flesh and shell of the animal, is
suffused with oil. Detailed analysis of this oil reveals that it is com
posed largely of polyunsaturated fatty acids of the linoleic type;
dodecanoic and much tetradecanoic acid are present. It has been
postulated that this oil is used largely for buoyancy, but this seems
unlikely—a leatherback has no more need of buoyancy than any
725
www.Ebook777.com
other sea turtle, and in fact probably less need, since its powerful
flippers allow it to rise to the surface whenever necessary, even
against a slight negative buoyancy. Moreover, all evidence points
to this being a deep-water species, to which marked buoyancy
would have a definite disadvantage. It is possible that, as in
certain whales, the oil has a function in lessening decompression
problems during rapid diving and re-surfacing.
The leatherback turtle used to be considered so rare that news
paper articles and learned papers were written every time a speci
men came to light; as late as 1961 it was estimated that there were
only about a thousand pairs in the world. However, we now know of
a number of good nesting beaches for the species, and I have
recently estimated the world population of mature females of this
species to be between 29,000 and 40,000. The nesting range of the
species can be summarized as follows:
In the Indian Ocean there is a leatherback beach of moderate
importance on the Tongaland coast of Natal, and leatherbacks also
nest on the coast of Ceylon. The best known Asiatic nesting ground
is located on 7.5 miles of beach in central Trengganu, Malaysia;
here several dozen individuals may nest on a good night. There are
as yet unevaluated nesting beaches in New Guinea and on the west
coast of Thailand, near and on the island of Pukhet. In the East
Pacific sporadic nesting occurs along the entire west coast of the
Americas, from Jalisco, Mexico, to Panama. Some pockets of
moderately concentrated nesting occur, e.g. at Ipala and at Mis-
maloya, Jalisco, and at Playa Manzanillo and Playa Cuyutlan,
Michoacan. The best is at the Playa de Bahia Chacahua, near
Punto Galera, Oaxaca, where up to 100 leatherbacks may nest
each night from September to December.
In West Africa some nesting occurs from Angola to Senegal,
but details are lacking. There are a few isolated nesting records
for Florida, and there is a definite leatherback ‘rookery’ at
Matina, Costa Rica. A few nest on the Santa Marta Peninsula,
Colombia, and in the Northwest District of Guyana. Bigisanti
Beach, Surinam, has several leatherbacks nesting each night
during the season. I, together with my colleagues Johan Schulz
and William Greenhood, had the honor of making known to the out
side world by far the best leatherback beach of all. On about ten
miles of beach in extreme western French Guiana leatherbacks
nest in enormous numbers—up to about 300 per night during the
peak months of May and June. In 1976 it appeared that both
Bigisanti and French Guiana beaches had been virtually deserted,
but in 1977 Jacques Fretey tagged 5800 nesting leatherbacks
in French Guiana. Many turtles now nest in Surinam.
Leatherbacks very rarely nest on islands (though there are
isolated records for Fiji, Jamaica, St. Kitts, and Nevis) and
726
Hatchling leatherback turtle, Dermochelys coriacea. Notice the distinct

plates on the shell and the large yolk sac.
727
www.Ebook777.com
eschew areas where coral or jagged rocks are present. This is
understandable, since the skin of the leatherback is so tender that
the animal may suffer multiple' minor lacerations just from
moving along a sandy beach, and I have seen nesting females
bleeding profusely from repeated contact with stumps and roots
that obstruct their path or their digging efforts on the beach.
The nesting behavior of the leatherback is well-known; indeed,
this is about the only aspect of the life history that is well known,
since the animals manage to keep themselves astonishingly well
hidden when at sea and only occasionally get caught, deliberately
or accidentally, by man. Nesting takes place by night, and an indi
vidual may nest several times (possibly as many as nine times in
some cases) in the course of a season, at intervals of, on average,
about ten days. This average internesting interval is distinctly
shorter than that of any other sea turtle. Relatively few data on the
intervals between nesting seasons are available, but it seems
probable that normally leatherbacks nest in alternate years, with
a small number nesting in successive years or at three-year inter
vals. The nesting season is typically from March to July in the
northern hemisphere and from October to February in the south
ern.
The nesting leatherback is a relatively imperturbable animal;
lights do not disturb them much, and they may be tagged as they
leave the sea without interrupting the nesting sequence. Moreover,
they nest even on very rainy nights, when most turtles of other
species avoid coming on shore. The trip up to the top of the beach
usually follows a sinusoidal track, the simplest explanation for
which is that it lessens the effective slope up which the ponderous
and (on land) clumsy animal has to climb. The sequence of events
preceding and following egg deposition follows essentially the pat
tern for all sea turtles as outlined earlier in this chapter. The
‘tears’ of a nesting leatherback are particularly viscous and often
form sand-covered ribbons several inches long hanging from each
eye. The body pit may be very shallow or may be so deep as to
lower the top of the turtle’s shell to the level of the undisturbed
sand. While moving on land, leatherbacks emit loud groans and
belches, as if the animal was not only in great pain but was also
suffering from severe indigestion. Another idiosyncrasy of the
nesting leatherback is its habit of turning a complete circle, or
even several circles, after completing the nest. The purpose of this
habit is obscure, but it may be concerned with re-orientation
toward the sea. I have seen some leatherback tracks in French
Guiana in which the animal had turned in about twenty complete
circles before re-entering the sea. A nesting leatherback is usually
on the shore for 1.5 to 2 hours.
728
Accounts of leatherbacks laying 300 eggs, or even 1000 eggs,

such as may be found in the literature, are fictitious. In fact the
leatherback lays rather few eggs for a sea turtle; in the Guianas
the average clutch numbers around 86 eggs, though in Tongaland,
South Africa, the average is about 106. The average in Trengganu
is 82.3, while in the East Pacific the average may be as low as 50.
These counts do not include the undersized, yolkless, and other
wise malformed eggs that are present in every nest of this species.
The number of these abnormal eggs is very variable, but it prob
ably averages 20 to 40% of the number of normal eggs. The eggs
are a little more than two inches in diameter and take 60 to 70 days
to hatch. The hatchlings are about 214” in carapace length and are
similar to the adults in appearance, though the fore flippers are
proportionately even longer and both shell and skin are covered
with small scales. These scales disappear within a couple of
months, though indistinct wrinkles mark the former scale boun
daries for some time later. The hatchling is marked with pure
white along the carapace ridges and the flipper margins, but this
pattern later changes to the more random spotting of the adult.
Baby leatherbacks frequently turn in ‘orientation circles’ as
they head for the sea, as do the adults. They are not easy to raise in
captivity, and indeed until recently the only baby leatherbacks
that had survived more than about two months under captive con
ditions were two raised by P.E.P. Deraniyagala in Ceylon in the
1930’s, one of which survived for 662 days and reached a carapace
length of about 16.5 inches and a weight of about 16 lbs. The other
died after 169 days. However, E. Birkenmeier of the Hassanal Bol-
kiah Aquarium in Brunei recently raised a leatherback to an age of
almost one year and a carapace length of 18.9”. This is thus a con
siderably better growth rate than was shown by Deraniyagala’s
specimen, though a poorer longevity. Recently the Miami Sea-
quarium raised several baby Florida leatherbacks to an age of
about eight months before they perished during a cold spell. Three
more hatchlings were obtained by the Seaquarium in July 1973,
and two of these animals were still alive in March 1974. Growth
rates have been most impressive; at the age of eight months the
turtles weighed about 35 lbs. each.* One of these grew from six to
fifteen pounds in only a month, and if growth continues in this ex
ponential fashion, mature size could be reached in three years or
less. These turtles were fed entirely upon jellyfish (Cassiopea), of
which they were capable of consuming about twice their own
weight daily. The great fluid intake associated with this diet
appears to be lost largely through the eyes, whose tremendously
copious secretion is evident as soon as the animals are lifted out of
* One weighed 6Jf lbs. at 22 months.
729
www.Ebook777.com
the water. Leatherbacks in the wild appear to feed largely upon
jellyfish also, together with some tunicates; captive specimens fed
upon fish die even more quickly than those not fed at all, apparent
ly from intestinal impaction. No adult has been kept for longer
than three weeks, with the exception of one which survived in the
aquarium in Casablanca, Morocco, for about six months, though
without feeding. This diet is of course well in keeping with the
weak, scissor-like jaws of the leatherback; but one wonders how
many jellyfish they have to eat to reach a weight of over 1000 lbs.!
Ingestion of jellyfish is assisted by an extensive arrangement of
flexible, backwardly-directed, pointed papillae which cover the
palate, throat, and esophagus of the leatherback. These are pre
sumably actively peristaltic in the living animal.
Little is known of leatherback migrations, but they are surely
extensive. There appears to be some sort of migration, possibly
from nesting grounds in New Guinea, down the east coast of Aus
tralia, and leatherbacks show up in the western Mediterranean
and off the Maritime Provinces of Canada more frequently than
chance drifting would suggest. However, those leatherbacks which
reach the shores of the British Isles are frequently dead on arrival.
The only long-distance recoveries of tagged individuals on record
are those of five females tagged while nesting in the Guianas on
Bigisanti Beach, Surinam, and on Silebache Beach, French
Guiana, one of which was recovered in Ghana, 6,800 kilometers
away, only a few months later; the others were caught off the
coast of South Carolina; near Freeport, Texas; near Highlands,
New Jersey; and in Campeche, Mexico. Presumably these indi
viduals wandered closer to land than is usual for the species, but
nevertheless the results are indicative of an enormously wide-
ranging species.
Leatherbacks intermediate in size between hatchlings and
adults are almost never found, and it is probable that these remain
at a great depth in the ocean, as do the adults except when in the
vicinity of a nesting beach. Support is lent to this supposition by the
leatherback’s total inability to cope with obstacles in its path; in
captivity, in water or on land, it bashes itself into the walls of its
container until it kills itself. The only place where such an animal
could survive is the open ocean. However, if this supposition of a
deep-water habitat is true, it raises the question of how the animal
respires and metabolizes in the low temperatures of the abyssal
ocean. The answer to the first question may be that, like other soft-
skinned turtles (such as musk turtles and snapping turtles), exten
sive oxygen exchange takes place through the skin. It is known
that leatherbacks have a muscular sphincter in the pulmonary ar
teries which is apparently capable of diverting blood flow from the
pulmonary to the systemic circulation. Perhaps when the animal
730
dives it simply cuts off all but the minimal blood flow to the lungs
and obtains enough oxygen for its needs through the extensive
superficial capillaries, which lie so near the surface on the under
side of the animal that the unpigmented areas are often strongly
suffused with pink. As far as metabolism and maintenance of high
activity at low temperatures is concerned, fragmentary evidence
suggests that the leatherback, like certain pelagic fish, can main
tain a body temperature considerably in excess of that of its sur
roundings. Recent studies by Wayne Frair, R.G. Ackman, and
Nicholas Mrosovsky have shown that a leatherback in a tank of
cold water can show a deep body temperature 18°C higher than
that of its surroundings. The animal is morphologically well
designed for the maintenance of such a substantial temperature
differential; the large size of the leatherback ensures that the sur-
face/volume ratio is small, and the barrel-shaped body would les
sen this ratio still more. Moreover the thick, cartilaginous shell is
almost the equivalent of a layer of blubber in insulating the animal
from its surroundings, and heat loss from the extremities is prob
ably minimized by a countercurrent heat exchange mechanism.
However, leatherbacks in the warm water near their nesting
beaches are only 2-3° warmer than their surroundings.
Clifford Pope, in his book The Reptile World, described the
leatherback as ‘the fly in the ointment of turtle classification.’ If
one may, perhaps incongruously, juxtapose a biblical quotation,
that ‘dead flies in the apothecary’s ointment give forth a stinking
savour,’ the conclusion of our labored syllogism is indeed true; a
dead leatherback rapidly develops a most characteristic and un
pleasant smell. This is caused by the decomposing oil, which is
present throughout the animal in great abundance and which is
very difficult to remove from anything with which it comes in con
tact. Nevertheless, this oil, when fresh, is supposed to have medici
nal properties in some areas, including in the American Virgin
Islands, and enormous prices may be paid for it. Otherwise the
leatherback is normally considered unfit for human consumption,
though a limited leatherback fishery exists on the coast of Peru. In
some places the oil is used for caulking small boats; for example,
about 10-15 leatherbacks are caught annually near Larak Island in
the Persian Gulf and are slaughtered for the sake of their oil.
Two subspecies of the leatherback have been described: Der-
mochelys coriacea coriacea in the Atlantic Ocean and D.c. schle-
geli in the Indian and Pacific Oceans. The name schlegeli, how
ever, is a nomen nudum, and the race is based simply on an illus
tration in Temminck and Schlegel’s Fauna Japonica. The head of
the animal in this illustration seems to be narrower than is normal
in Atlantic leatherbacks, but there is little doubt that the drawing
is simply inaccurate; eight leatherback skulls I collected in Peru
751
www.Ebook777.com
were no narrower, on average, than a series of twelve from French
Guiana, though they were more variable. It has also been sugges
ted that coriacea is darker and less spotted than schlegeli; but the
coloration is extremely variable even within the French Guiana
nesting population, which includes both heavily spotted and almost
uniformly black individuals. There is some indication, however,
that East Pacific leatherbacks mature at a smaller size than those
from the Atlantic; seven carapaces I measured in Peru were all
less than four feet long, and a skull I found on a nesting beach in
Guerrero was distinctly smaller than any mature Atlantic leather
back skulls I have seen.
The leatherback would seem to have a moderately good survi
val outlook, since there are few places where the adult turtles are
killed, though excessive egg collection takes place on several of
the nesting grounds, including the important Costa Rican and
Malaysian beaches. However, egg collection is prohibited in
French Guiana, where probably nearly half the leatherbacks in
the world nest, and very little illicit collection takes place. The
adults would seem to have few natural enemies at sea, but one
finds that a high proportion of nesting females are mutilated, pre
sumably by sharks. On Bigisanti Beach, Surinam, a few nesting
leatherbacks are killed by jaguars, and parts of a leatherback
have been recovered from the stomachs of three killer whales
caught near St. Vincent, in the West Indies.
Leatherbacks are currently being studied on the nesting beach
in Tongaland by George Hughes and his co-workers; in Trinidad
by Peter Bacon and the Trinidad field-naturalists club; in Treng-
ganu by Tho Yew Pong and his co-workers; in Surinam by Johan
Schulz and the Surinam Forest Service; and in French Guiana by
myself and co-workers.
BACKGROUND READING
CHELONIA DEPRESSA:
Bustard, H.R. 1969. The flatback. Animals, Dec. 1969, pp. 356-
357.
__________ 1972. Australian sea turtles. Their Natural History
and Conservation. London, 220 pp.
__________and C. Limpus. 1969. Observations on the flatback
turtle Chelonia depressa Garman. Herpetologica 25(1):
29-34.
Cogger, H.G. and D.A. Lindner. 1969. Marine turtles in
Northern Australia. Aust. Zool. 15: 150-159.
732
Fry, D.B. 1913. On the status of Chelonia depressa Garman.

Rec. Aust. Mus. 10: 159-185.
Garman, S. 1880. On certain species of the Chelonioidae. Bull.
Mus. Comp. Zool., 6: 123-126.
Limpus, C. 1971. The flatback turtle, Chelonia depressa Gar
man in Southeast Queensland, Australia. Herpetologica 27:
431-446.
McCulloch, A.R. 1908. A new genus and species of turtle from
North Australia. Rec. Austral. Mus., 7: 126-128.
Williams, E.E., A.G.C. Grandison and A.F. Carr. 1967. Chelonia
depressa Garman re-investigated. Breviora, Mus. Comp.
Zool. 271: 1-15.
CHELONIA MYDAS:
Alvarez del Toro, M. 1960. Reptiles de Chiapas. Tuxtla Gutier
rez, Chiapas, Mexico, Institute de Zoologia del Estado, pp.
7-204.
Appa Rao, T. and S. Dutt. 1965. Chemical composition of the
flesh of the turtle Chelone mydas (Boulenger). Curr. Sci.,
34: 695.
Arabian American Oil Company, Aramco Handbook. Zonen-
Haarlem, J. Enschede, 279 pp.
Babcock, H.L. 1937. The sea turtles of the Bermuda Islands,
with a survey of the present state of the turtle fishing indus
try. Proc. Zool. Soc. London (A), 107: 595-601.
Balasingham, E. 1965. Conservation of green turtles (Chelonia
mydas) in Malaya. Malay Nat. J. 19: 235-6.
Balazs, G.H. 1975. Green turtle’s uncertain future. Defenders,
50(6): 521-523.
__________ 1976. Green turtle migrations in the Hawaiian Arch
ipelago. Biological Conservation, 9: 125-140.
__________ andE. Ross. 1974. Observations of the basking habit
in the captive juvenile Pacific green turtle. Copeia, 1974(2):
542-544.
Banks, E. 1937. The breeding of the edible turtle (Chelone my
das)i. Sarawak Mus. J., 4: 523-532.
Barth, R. 1962. Observanoes sobre a grande tartaruga marinha,
Chelonia mydas L., feitas na ilha de Trindade. An. Acad.
Bras. Cienc., 34: 405-409.
Baur, G. 1890. The genera of the Cheloniidae. Am. Nat. 24: 486-
487.
Berkson, H. 1966. Physiological adjustments to prolonged
diving in the Pacific green turtle (Chelonia mydas agassi-
zii). Comp. Biochem. Physiol., 18(1): 101-119.
__________. 1967. Physiological adjustments to deep diving in
the Pacific green turtle (Chelonia mydas agassizii). Comp.
Biochem. Physiol., 21(3): 507-524.
733
www.Ebook777.com
Booth, J. and J.A. Peters. 1972. Behavioural studies on the
green turtle (Chelonia mydas) in the sea. Animal Behaviour,
20(4): 808-812.
Boyer, D.R. 1965. Ecology of the basking habit in turtles. Ecol
ogy, 46: 99-118.
Brongersma, L.D. 1961. Notes upon some sea turtles. Zool.
Verh., Leiden, 51: 1-46.
__________ 1968. Notes upon some turtles from Surinam. Proc.
K. Ned. Akad. Wet. (C), 71: 114-127.
Bustard, H.R. 1967. Mechanism of nocturnal emergence from
the nest in green turtle hatchlings. Nature, London, 214: 317.
__________ 1970. The adaptive significance of coloration in
hatchling green sea turtles. Herpetologica, 26: 224-227.
__________ 1972. Australian sea turtles. London and Sydney, 220
pp.
__________ 1973. Kay’s turtles. London and Sydney, 128 pp.
__________and P. Greenham. 1969. Nesting behaviour of the
green sea turtle on a Great Barrier Reef Island. Herpeto
logica, 25: 93-102.
Caldwell, D.K. 1962a. Sea turtles in Baja Californian waters
(with special reference to those of the Gulf of California),
and the description of a new subspecies of northeastern
Pacific green turtle. Contrib. Sci. Los Angeles County Mus.,
(61): 1-31.
__________ 1962b. Carapace length-body weight relationship
and size and sex ratio of the northeastern Pacific green sea
turtle, Chelonia mydas carrinegra. Contrib. Sci. Los Angeles
County Mus., (62): 3-10.
__________. 1969. Hatchling green sea turtles, Chelonia mydas,
at sea in the northeastern Pacific Ocean. Bull. South. Calif.
Acad. Sci., (68): 113-114.
Carr, A.F. 1942. Notes on sea turtles. Proc. New England Zool.
Club, 21: 1-16.
__________. 1952. Handbook of Turtles. Ithaca, N.Y. Cornell
Univ. Press, 542 pp.
__________. 1956. The windward road. New York, Alfred Knopf
Inc. 258 pp.
__________ 1961. Pacific turtle problems. Nat. Hist., 70: 64-71.
__________. 1962a. Orientation problems in the high seas travel
and terrestrial movements of marine turtles. Am. Sci., 50:
359-374.
__________. 1962b. Guideposts of animal navigation. Am. Inst.
Biol. Sci., Boston, D.C. Heath Co., B.S.C.S. No. 1.
__________ 1965. The navigation of the green turtle. Sci. Am.,
212(5): 79-86.
734
__________ 1967a. Adaptive aspects of the scheduled travel of

Chelonia. In Animal Orientation and navigation, edited by
R.M. Storm, Corwallis, Oregon State University Press, pp.
35-52.
__________. 1967b. So Excellent a Fishe. New York, Natural His
tory Press, 248 pp.
__________. 1970. Green sea turtles in peril. Natl. Parks Conserv.
Mag., 44: 19-24.
__________-1975. The Ascension Island green turtle colony. Co-
peia, 1975(3): 547-555.
__________and D.K. Caldwell. 1956. The ecology and migrations
of sea turtles. 1. Results of field work in Florida, 1955. Am.
Mus. Novit., 1793: 1-23.
__________ and M.H. Carr. 1970. Modulated reproductive per
iodicity in Chelonia. Ecology, 51(2): 335-337.
_________ and__________ . 1972. Site fixity in the Caribbean green
turtle. Ecology, 53(3): 425-429.
__________and P.J. Coleman. 1974. Seafloor spreading theory
and the odyssey of the green turtle. Nature, 249(5453):
128-130.
__________and L. Giovannoli. 1957. The ecology and migrations
of sea turtles. 2. Results of field work in Costa Rica, 1955.
Am. Mus. Novit., 1835: 1-32.
__________and D. Goodman. 1970. E cologic implications of size
and growth in Chelonia. Copeia, 1970: 783-786.
__________and H. Hirth. 1961. Social facilitation in green turtle
siblings. Anim. Behav., 9(1-2): 68-70.
__________ 1962. The ecology and migrations of sea turtles, 5.
Comparative features of the isolated green turtle colonies.
Am. Mus. Novit., 2091: 1-42.
__________and L. Ogren. 1960. The ecology and migrations of
sea turtles. 4. The green turtle in the Caribbean Sea. Bull.
Amer. Mus. Nat. Hist., 121: 1-48.
__________, P. Ross, and S. Carr. 1974. Intemesting behavior of
the green turtle, Chelonia mydas, at a mid-ocean island
breeding ground. Copeia, 1974(3): 703-706.
__________and S. Stancyk. 1975. Observations on the ecology
and survival outlook of the hawksbill turtle. Biological Con
servation, 9: 161-172.
__________andD. Sweat. 1969. Long-range recovery of a tagged
yearling Chelonia on the east coast of North America. Biol.
Conserv., 1: 341-342.
de Silva, G.S. 1969. Statement on marine turtles in the State of
Sabah. IUCN Publ. New Ser. Suppl. Pap., 20: 75-79.
Colombo, 412 pp.
735
www.Ebook777.com
Domantay, J.s. 1953. The turtle fisheries of the Turtle Islands.
Bull. Fish. Soc. Philipp., 3-4, 3-27.
__________ 1968. Notes on the development and breeding habits
of Chelonia mydas. Acta Manilana (A), 4: 87-109.
Donoso-Barros, R. 1966. Reptiles de Chile. Santiago, Univ. of
Chile. 458 pp.
Duncan, D.D. 1944. Capturing giant turtles in the Caribbean.
Natl. Geogr. Mag., 84: 177-190.
Ehrenfeld, D.W. 1968. The role of vision in the sea-finding orien
tation of the green turtle (Chelonia mydas). 2. Orientation
mechanism and range of spectral sensitivity. Anim. Behav.,
16: 281-287.
__________ 1974. Conserving the edible sea turtle: can maricul-
ture help? American Scientist, 62: 23-31.
Ehrenfeld, D.W. and A. Koch. 1967. Visual accommodation in
the green turtle. Science, Wash., 155(3764): 827-828.
FAO/UN. 1968. Report to the Governments of the People’s Re
public of Southern Yemen and the Seychelles Islands on the
green turtle resource of South Arabia, and the status of the
green turtle in the Seychelles Islands. Based on the work of
Dr. H. Hirth, FAO/TA Marine Turtle Biologist. Rep. FAO/
UNDP(TA), 2467: 59 pp.
Felger, R.S., K. Cliffton, and P.J. Regal. 1976. Winter dormancy
in sea turtles: independent discovery and exploitation in the
Gulf of California by two local cultures. Science, 191: 283-285.
Ferreira, M.M. 1968. Sobre a alimentacao de aruana, Chelonia
mydas Linnaeus, ao longo da costa do estado do Ceara. Arq.
Estac. Biol. Mar. Univ. Fed. Ceara, 8: 83-86.
Frazier, J. 1971. Observations on sea turtles at Aldabra Atoll.
Phil. Trans. Roy. Soc. London, B. 260: 373-410.
Harrisson, T. 1950. The Sarawak Turtle Islands’ “Semah.” J.
Malay Branch, R. Asiatic Soc., 23: 105-126.
__________ 1951 to 1967. Eighteen papers on green turtles in
Borneo in the Sarawak Museum Journal.
__________ 1969. The marine turtle situation in Sarawak. IUCN
Publ. New. Ser. Suppl. Pap., (20): 70-74.
Hendrickson, J.R. 1958. The green sea turtle, Chelonia mydas
(Linn.), in Malaya and Sarawak. Proc. Zool. Soc. Lond., 130:
455-535.
Hill, R.L. and D.J. Green. 1971. Surinam turtle notes — 2: In
vestigation of the damage by the crab Ocypode quadrata to
the eggs of the green turtle, Chelonia mydas. STINASU,'
Surinam: pp. 11-13.
Hirth, H.F. 1971. Synopsis of biological data on the Green Tur
tle. FAO Fisheries Synopsis No. 85, pp. 1:1 to 8:19.
736
Hirth, H.F. and A.F. Carr. 1970. The green turtle in the Gulf of
Aden and the Seychelles Islands. Verh. K. Ned. Akad. Wet.
(Afd. Nat. Tweede Sect.), 58: 1-44.
Koch, A.L., A.F. Carr and D.W. Ehrenfeld. 1969. The problem of
open-sea navigation: the migration of the green turtle to
Ascension Island. J. Theor. Biol., 22: 163-179.
McCoy, M.A. 1974. Man and turtle in the Central Carolines.
Micronesica, 10(2): 207-221.
Marquez, R. and T. Doi. 1973. Ensayo teorico sobre el analisis
de la poblacion de tortuga prieta, Chelonia mydas carrin-
egra Caldwell, en aguas del Golfo de California, Mexico.
Bull. Tokai Regional Fisheries Research Lab., No. 73: 1-22.
Mellem, I.M. 1925. Marine turtles sleep on Hawaiian sands.
Bull. N.Y. Zool. Soc., 28: 160-161.
Minton, S.A. 1966. A contribution to the herpetology of West
Pakistan. Bull. Am. Mus. Nat. Hist., 134: 27-184.
Moorehouse, F.W. 1933. Notes on the green turtle (Chelonia my
das). Rep. Great Barrier Reef Comm., 4(1): 1-22.
Mrosovsky, N. 1970. The influence of the sun’s position and
elevated cues on the orientation of hatchling sea turtles.
Animal Behavior, 18: 648-651.
__________ 1972. The water-finding ability of sea turtles. Brain,
Behavior and Evolution, 5(2/3): 202-225.
_________ and A.F. Carr. 1967. Preferences for light of short
wavelengths in hatchling green sea turtles, Chelonia mydas,
tested on their natural nesting beaches. Behaviour, 28(3/4):
217-231.
_________ and S.J. Shettleworth. 1968. Wavelength preferences
and brightness cues in the water-finding behaviour of sea
turtles. Behaviour, 32: 211-257.
Parsons, J.J. 1962. The green turtle and man. Gainesville, Univ.
of Florida Press, 126 pp.
Peters, J.A. 1956-57. The eggs (turtle) and I. Biologist, 39: 21-24.
Pritchard, P.C.H. 1969. Sea turtles of the Guianas. Bull. Fla.
State Mus., Biol. Sci., 13: 85-140.
__________ 1971. Galapagos Sea Turtles—Preliminary Findings.
J. Herpetology, 5: 1-2 pp. 1-9.
__________ 1973. International Migrations of South American
sea turtles (Cheloniidae and Dermochelidae). Animal
Behav., 21(1): 18-27.
Rainey, W.E. 1971. Reconnaissance of the green turtle, Chelo
nia mydas, nesting aggregation at Aves Island, Lesser Antil
les. ms.
------------ and P.C.H. Pritchard. 1972. Distribution and man-
737
www.Ebook777.com
agement of Caribbean sea turtles. Contrib. No. 105, Virgin
Islands Ecological Research Station, 1-21.
Rebel, T.P. 1974. Sea turtles and the turtle industry of the West
Indies, Florida, and the Gulf of Mexico. Univ. of Miami
Press. 250 pp.
Schmidt, J. 1916. Marking experiments with turtles in the
Danish West Indies. Medd. Komm. Harvunders. (Ser.
Fisk.), 5: 1-26.
Schulz, J.P. 1968. Zeeschildpadden, Deel II: Zeeschildpadden
in Suriname. Dienst Lands Bosbeheer, Suriname; 103 pp.
Shah, R.V. 1962. A comparative study of the respiratory
muscles in Chelonia. Breviora, 161: 1-16.
Travis, W. 1967. The voice of the turtle. London, George Allen
and Unwin, 203 pp.
Urban, E.K. 1970. Nesting of the green turtle (Chelonia mydas)
in the Dahlak Archipelago, Ethiopia. Copeia 1970(2): 393-
394.
caise. Init. Afr., (15): 354 pp.
Witham, R. and A.F. Carr. 1968. Returns of pen-reared green
turtles. Quart. J. Fla. Acad. Sci., 31: 49-50.
LEPIDOCHELYS KEMPI AND LEPIDOCHELYS OLIVACEA:
Aguayo, C.G. 1953. La tortuga bastarda (Lepidochelys olivacea
kempi) en Cuba. Mem. Soc. Cubana de Hist. Nat., 21(2):
211-219.
Alvarez del Toro, M. 1960. Los Reptiles de Chiapas. Inst. Zool.
del Estado, Chiapas, Mexico: 204 pp.
Anon. 1966. Marcado de tortugas en el Pacifico Mexicano. Inst.
Nac. Inv. Biol. Pes. Bol. Programa Nacional de Marcado de
Tortugas Marinas 1(4): 1-8.
Baur, G. 1890. The genera of the Cheloniidae. Amer. Nat., 24:
486-487.
Bleakney, S. 1965. Reports of marine turtles from New England
and Eastern Canada. Canadian Field-Naturalist, 79(2): 120-
128.
Verh. Mus. Leiden, 51: 1-46.
__________ 1972. European Atlantic turtles. Zool. Verh. Mus.
Leiden, 121: 1-318.
Caldwell, D.K. 1966. A nesting report on the American ridley.
Int. Turt. Tort. Soc. J., 1: 10-13, 30.
------------ and R.S. Casebeer. 1964. A note on the nesting of the
eastern Pacific ridley sea turtle, Lepidochelys olivacea.
Herpetologica, 20(3): 213.
738
__________, W.F. Rathjen and B.C.C. Hsu. 1969. Surinam ridleys

at sea. Int. Turt. Tort. Soc. J. 3: 4-5, 23.
Club, 21: 1-16.
__________. 1948. Sea turtles on a tropical island. Fauna, 10:
50-55.
__________. 1952. Handbook of turtles. Comstock Publ. Assoc.,
Cornell Univ. Press. 542 pp.
__________. 1956. The Windward Road. Alfred Knopf, N.Y.
258 pp.
__________ 1957. Notes on the zoogeography of the Atlantic sea
turtles of the genus Lepidochelys. Rev. Biol. Trop. 5(1): 45-
61.
__________ 1961. The ridley mystery today. Animal Kingdom, 64
(1): 7-12.
__________ 1963. Panspecific reproductive convergence in Lepi
dochelys kempi. Ergebn. Biol., 26: 298-303.
__________ 1967. So Excellent a Flshe. Nat. Hist. Press, N.Y.
248 pp.
_____ and D.K. Caldwell. 1956. The ecology and migrations
of sea turtles, 1. Results of field work in Florida, 1955. Amer.
Mus. Novitates, 1793: 1-23.
____________________. 1958. The problem of the Atlantic ridley
turtle (Lepidochelys kempi) in 1958. Rev. Biol. Trop. 6(2):
245-262.
Chavez, H. 1968. Marcado y recaptura de individuos de tortuga
lora, Lepidochelys kempi (Garman). Inst. Nac. Inv. Biol.
Pes. 19: 1-28.
__________, M. Contreras and E. Hernandez. 1967. Aspectos bio-
logicos y proteccion de la tortuga lora, Lepidochelys kempi
(Garman), en la costa de Tamaulipas, Mexico. Inst. Nac.
Inv. Biol. Pes. 17: 1-40.
Colombo. 412 pp.
Dodge, E.S. 1944. Status of the ridley turtle in Massachusetts
waters. Copeia (2): 120-121.
Donoso-Barros, R. 1964. Nota sobre Lepidochelys kempi en las
costas de Cumana. Lagena, 2: 20-21.
Eder, H.M. 1969. Turtling in coastal Oaxaca. Pacific Discovery,
22(1): 10-15.
Flores, C. 1966. Nuevos registros de Lepidochelys kempi (Gar
man) en la costa oriental de Venezuela. Lagena, 12: 11-14.
Hardy, J.D. Jr. 1962. Comments on the Atlantic ridley turtle,
Lepidochelys olivacea kempi, in the Chesapeake Bay. Ches
apeake Science, 3(3): 217-220.
739
www.Ebook777.com
Hay, O.P. 1908. On three existing species of sea-turtles, one of
them (Caretta remivaga) new. Proc. U.S. Nat. Mus., 34:
183-198.
Hildebrand, H.H. 1963. Hallazgo del area de anidacion de la tor-
tuga marina ‘lora’, Lepidochelys kempi (Garman), en la
costa occidental delGolfode Mexico (Rept., Chel.). Ciencia,
Mexico, 22(4): 105-112.
Hill, R.L. 1971a. Surinam turtle notes — 1: Polymorphism of

coastal and vertebral laminae in the sea turtle Lepidochelys
olivacea. STINASU, Surinam: pp. 3-9.
__________ 1971b. Surinam turtle notes — 3: The effect of rup
turing eggs in sea turtle nests on the hatchling emergence
percentage. STINASU, Surinam: pp 14-16.
Houck, W.J. and J.G. Joseph. 1958. A northern record for the
Pacific ridley, Lepidochelys olivacea. Copeia, 1958(3): 219-
220.
Hughes, D.A. and J.D. Richard. 1974. The nesting of the Pacific
ridley turtle Lepidochelys olivacea on Playa Nancite, Costa
Rica. Marine Biology, 24 : 97-107.
Loveridge, A. andE.E. Williams. 1957. Revision of the African
Comp. Zool., 115(6): 163-557.
Marquez, R. In press. Las tortugas marinas de Mexico. Inst.
Nac. Inv. Biol. Pes., Mexico.
Mowbray, L.S. and D.K. Caldwell. 1958. First record of the rid
ley turtle from Bermuda, with notes on other sea turtles and
the turtle fishery in the islands. Copeia, (2): 147-148.
Oliver, J.A. 1946. An aggregation of Pacific sea turtles. Copeia,
(2): 103.
Pope, C.H. 1935. The reptiles of China. Nat. Hist, of Central
Asia, 10. Am. Mus. Nat. Hist. 604 pp.
Pritchard, P.C.H. 1966. Occurrence of mesoplastra in a Crypto-
diran turtle, Lepidochelys olivacea. Nature, 210: 652.
------------- 1967. To find the ridley. Int. Turt. Tort. Soc. J. 1(4):
30-35, 48.
------------- 1969a. Sea turtles of the Guianas. Bull. Florida State
Mus., 13(2): 85-140.
------------- 1969b. The survival status of ridley sea-turtles in
American waters. Biol. Conserv. 2(1): 13-17.
__________- 1973. International migrations of South American
Behav. 21(1): 18-27.
------------ and R. Marquez. 1974. Kemp’s ridley turtle, or the
Atlantic ridley Lepidochelys kempi. IUCN Monograph, No.
2.
740
Schmidt, K.P. and E.R. Dunn. 1917. Notes on Colpochelys

kempi Garman. Copeia, No. 44: 50-52.
Zahl, P.A. 1973. One strange night on turtle beach. Nat. Geogr.
Mag., Oct. 1973, pp. 570-581.
DERMOCHELYS CORIACEA:
Allen, E.R. and W.T. Neill. 1957. Another record of the Atlantic
leatherback, Dermochelys coriacea coriacea, nesting on the
Florida coast. Copeia, 1957 (2): 143-144.
Bacon, P.R. 1969. Report on the Trinidad sea turtle conserva
tion project. Annual Report of the Trinidad Field Naturalists
Club for 1969, pp. 18-35.
__________. 1970. Studies on the leatherback turtle, Dermo
chelys coriacea (L.), in Trinidad, West Indies. Biol. Con-
serv. 2(3): 213-217.
__________. 1973. The orientation circle in the beach ascent
crawl of the leatherback turtle, Dermochelys coriacea, in
Trinidad. Herpetologica, 29(4): 343-348.
Balasingham, E. 1967. The ecology and conservation of the
leathery turtle, Dermochelys coriacea (Linn.) in Malaya.
Proc. 11th. Pacific Science Congress, Tokyo: 37-43.
Brongersma, L.D. 1967. British Turtles. Brit. Mus. (Nat. Hist.),
i-viii, 1-22.
__________. 1968a. The great sea serpent and the leathery turtle,
I. Proc. Kon. Ned. Akad. Wetensch. Amsterdam, C 71: 209-
228.
__________. 1968b. Notes upon some sea turtles from the Canary
Islands and from Madeira. Proc. Kon. Ned. Akad. Wetensch.
Amsterdam, C 71: 128-136.
__________. 1969. Miscellaneous notes on turtles, II A. ibid, C 72;
76-102.
__________. 1972. European Atlantic Turtles. Zool. Ver. Mus.
Leiden, 121: 1-318.
Caldwell, D.K. 1959. On the status of the Atlantic leatherback
turtle, Dermochelys coriacea coriacea, as a visitant to
Florida nesting beaches, with natural history notes. Quart.
J. Fla. Acad. Sci. (1958), 21(3): 285-291.
__________and M. Caldwell. 1969. Addition of the leatherback
sea turtle to the known prey of the killer whale Orcinus orca.
J. Mammalogy, 50(3): 636.
Carr, A.F. 1952. Handbook of turtles. Comstock Publ. Assoc.,
__________and L. Ogren. 1959. The ecology and migrations of
sea turtles, 3: Dermochelys in Costa Rica. Amer. Mus. Novi-
tates No. 158: 1-29.
741
www.Ebook777.com
Deraniyagala, P.E.P. 1939. The tetrapod reptiles of Ceylon,
Vol. 1 Colombo. 412 pp.
Dunlap, C.E. 1955. Notes on the visceral anatomy of the giant
leatherback turtle (Dermochelys coriacea Linnaeus). Bull.
Tulane Med. Fac. 14(2): 55-69.
Fitter, R.S.R. 1961. The leathery turtle or luth. Oryx, 6(2): 116-
125.
Ford, J. 1879. The leather turtle. Amer. Nat. 13(10): 633-637.
Goeldi, E.A. 1906. Chelonios do Brasil. Bol. Mus. Goeldi do Hist.
Nat. Ethnographica (Para), 4: 699-756.
Hardy, J.D. 1969. Records of the leatherback turtle, Dermo
chelys coriacea coriacea (Linnaeus), from the Chesapeake
Bay. Bull. Maryland Herp. Soc. 5(3): 92-96.
Heldt, H. 1933. La tortue luth, Sphargis coriacea (L.). Captures
faites sur les cotes tunisiennes (1930-1933). Contribution ana-
tomique et biologique de l’espece. Ann. Stat. Oceanogr.
Salammbo, 8: 1-40, 17 figs.
Hughes, G.R. 1969. Report to the Survival Service Commission
on marine turtles in Southern Africa. In Marine Turtles:
Proceedings of the Working Meeting of Marine Turtle Spe
cialists organized by IUCN at Morges, Switzerland, 10-13
March 1969: 56-66.
__________ 1970. Further studies on marine turtles in Tonga-
land. Ill Lammergeyer, 12, 7-25.
__________ 1970. Further studies on marine turtles in Tonga-
land. IV Lammergeyer, 12, 26-36.
__________ 1971. The marine turtles of Tongaland. V Lammer
geyer, 13, 7-24.
__________ 1972. The marine turtles of Tongaland. 6 Lammer
geyer, 15, 15-26.
__________, A. J. Bass and M.T. Mentis. 1967. Further studies on
marine turtles in Tongaland. Lammergeyer, 7: 5-54.
Leary, T.H. 1957. A schooling of leatherback turtles, Dermo
chelys coriacea, on the Texas coast. Copeia, 1957: 232.
Comp. Zool. 115(6): 163-557.
McAskie, I.B. and C.R. Forrester. 1962. Pacific leatherback
turtles (Dermochelys) of the coast of British Columbia.
Copeia, 1962: 646.
Mrosovsky, N. and S.J. Shettleworth. 1975. On the orientation

circle of the leatherback turtle, Dermochelys coricct,a. Ani
mal Behaviour, 23(3): 568-591.
Nishimura, S. 1964. Considerations on the migration of the
leatherback turtle, Dermochelys coriacea, in the Japanese
742
and adjacent waters. Publ. Seto Marine Biol. Lab. 12(2):

61-73.
Pritchard, P.C.H. 1969. Sea turtles of the Guianas. Bull. Florida
State Mus. 13(2): 85-140.
__________. 1971. The leatherback or leathery turtle. IUCN
Monograph No. 1, 39 pp.
__________. 1973. International migrations of South American
Behav. 21: 18-27.
Schulz, J.P. 1968. Zeeschildpadden, Deel II: Zeeschildpadden in
Suriname. 2nd. Ed. Dienst Lands Bosbeheer, Paramaribo:
1-103.
_________ . 1970. Nesting beaches of sea turtles in west French
Guiana. Koninkl. Nederl. Akademie van Wetenschappen,
Amsterdam. Ser. C, Proc., 74(4): 396-404.
caise. Dakar. Init. Afr. 15: 1-345.
Willgohs, J.F. 1957. Occurrence of the leather turtle in the
Northern North Sea and off Western Norway. Nature, Lon
don, 179: 163-164.
Yerger, R.W. 1965. The leatherback turtle on the Gulf Coast of
Florida. Copeia, 1965: 364-366.
ERETMOCHELYS IMBRICATA:
Bustard, H.R. 1972. Australian sea turtles. Collins, London and
Sydney. 220 pp.
Caldwell, D.W. 1962. Growth measurements of young Atlantic
sea turtles in temperate waters. Contr. in Sci. Los Angeles
Co. Mus., 50: 1-8.
Carr, A.F. 1952. Handbook of turtles. Comstock Publ. Assoc.,
__________ 1956. The Windward Road. A. Knopf, New York.
258 pp.
__________ 1967. So Excellent a Fishe. Nat. Hist. Press, New
York. 248 pp.
__________, H. Hirth and L. Ogren. 1966. The ecology and migra
tions of sea turtles, 6. The hawksbill turtle in the Caribbean
Sea. Am. Mus. Novitates 2248, 1-29.
__________. 1972. Great reptiles, great enigmas. Audubon, 74(2):
24-35.
Colombo. 412 pp.
Frazier, J. 1971. Observations on sea turtles at Aldabra Atoll.
Phil. Trans. Roy. Soc. London B. 260, 373-410.
Hirth, H.F. 1962. Cloacal temperatures of the green and hawks
bill sea turtles. Copeia 1962: 647-648.
743
www.Ebook777.com
__________and A.F. Carr. 1970. The green turtle in the Gulf of
Aden and the Seychelles Islands. Ver. Kon. Nederlandse
Akad. Wetenschap. Nat. 58: 1-44.
Homell, J. 1927. The turtle fisheries of the Seychelles Islands.
London, H.M. Stationery Office, pp. 1-55.
Hutchison, A.D. 1969. Shipwreck search on the Caribbean
Banks. Explorers Journal, March 1969: 41-49.
Kinunen, W. and P. Walczak. Persian Gulf Sea Turtle Nesting
Surveys. Ms. submitted to Director, Iran Game and Fish
Department, 1971.
Lewis, C.B. 1940. The Cayman Islands and Marine Turtle. Bull.
Inst. Jamaica, 2 (Appendix): 56-65.
Tortoises and Turtles of the Suborder Cryptodira. Bull. Mus.
Comp. Zool. 115(6): 163-557.
Nietschmann, B. Unpublished ms. The exploitation of hawksbill
turtles, East Coast of Nicaragua: results and recommenda
tions from a preliminary study. 10 pp.
Nishimura, S. and T. Yasuda. 1967. Records of the hawksbill
turtle, Eretmochelys imbricata (Linne), in the Japan Sea.
Publ. Seto Marine Biol. Lab, XV(4): 297-302.
Pritchard, P.C.H. 1969. Sea turtles of the Guianas. Bull. Florida
State Mus., 13(2): 85-140.
Rainey, W.E. and P.C.H. Pritchard. 1972. Distribution and
management of Caribbean sea turtles. Contrib. No. 105,
Virgin Islands Ecological Research Station, 1-21.
Schmidt, J. 1916. Marking experiments with turtles in the
Danish West Indies. Meddelelser Kommissionen Havunder-
sogelser, Ser. Fiskeri 5: 1-26.
Steinbeck, J. 1962. The log from the Sea of Cortez. Viking Com
pass Books, 282 pp.
Suwelo, I.S. 1973. Notes on turtle ranching at Tidung Island.
Universitas Nasional, Jakarta. Pp. 1-3.
caise. I.F.A.N., Dakar. 354 pp.
CARETTA CARETTA:
Verh. Leiden, 51(2): 1-46.
__________ 1968a. Notes upon some sea turtles from Surinam.
Proc. Kon. Ned. Akad. Wet., Amsterdam, C 71, 2: 114-127.
__________ 1968b. Notes upon some sea turtles from the Canary
Islands and from Madeira. Proc. Kon. Ned. Akad. Wet.,
Amsterdam. C 71, 2: 128-136.
__________ 1972. European Atlantic turtles. Zool. Verh. Mus.
Leiden, 121: 1-318.
744
Bustard, H.R. 1972. Australian sea turtles. Collins, London and

Sydney. 220 pp.
__________, P.M. Greenham and C. Limpus. 1971. Nesting be
haviour of loggerhead and flatback turtles in Queensland,
Australia. Proc. Kon. Ned. Akad. Sci., C.
__________and C. Limpus. 1970. First international recapture of
an Australian tagged loggerhead. Herpetologica 26: 358-359.
____________________ 1971. Loggerhead turtle movements. Brit.
J. Herpetology. 4: 228-230.
__________, K. Simkiss and N.K. Jenkins. 1969. Some analysis of
artificially incubated eggs and hatchlings of green and
loggerhead sea turtles. J. Zool., London 158: 311-315.
Caldwell, D.K. 1959. The loggerhead turtles of Cape Romain,
South Carolina. Bull. Fla. State Mus., Biol. Sci., 4(10): 319-
348.
__________. 1962. Comments on the nesting behaviour of Atlantic
loggerhead sea turtles, based primarily on tagging returns.
Quart. J. Florida Acad. Sci. 25: 287-302.
__________. 1968. Baby loggerhead turtles associated with Sar-
gassum weed. Quart. J. Florida Acad. Sci. 31: 271-272.
__________, F.H. Berry, A.F. Carr and R.A. Ragotzkie. 1959.
Multiple and group nesting by the Atlantic loggerhead tur
tle. Bull. Fla. State Mus., Biol. Sci., 4(10): 309-318.
__________, A.F. Carr and T.R. Hellier. 1955. Natural history
notes on the Atlantic loggerhead turtle, Caretta caretta
caretta. Quart. J. Florida Acad. Sci. 18: 292-302.
__________, A.F. Carr and L. Ogren. 1959. Nesting and migration
of the Atlantic loggerhead turtle. Bull. Florida State Mus.,
Biol. Sci., 4(10): 295-308.
Club 21: 1-16.
__________. 1952. Handbook of turtles. Comstock Publ. Associ
ates, Cornell Univ. Press. 542 pp.
__________. 1956. The Windward Road. Alfred A. Knopf, Inc.,
New York. 258 pp.
__________. 1967. So Excellent a Fishe. Nat. Hist. Press, N.Y.
248 pp.
Deraniyagala, P.E.P. 1933. The loggerhead turtles (Caretti-
dae) of Ceylon. Ceylon J. Sci. (B), 18: 61-72.
__________. 1939. The tetrapod reptiles of Ceylon. Vol. 1. Colom
bo. 412 pp.
__________. 1952. The loggerhead turtles (Carettinae) of Europe.
Herpetologica 8: 57-58.
Gallagher, R.M., M.L. Hollinger, R.M. Ingle, and C.R. Futch.
1972. Marine turtle nesting on Hutchinson Island, Florida in
1971. Fla. Dept, of Nat. Res., Special Scientific Report No.
37, 11 pp.
745
www.Ebook777.com
Hughes, G.R. 1967. Marine turtle hatchlings of Tongaland. Afri
can Wild Life 23(1): 5-19.
_________ . 1970a. Further studies on marine turtles in Tonga
land, III. Lammergeyer 12: 7-25.
__________ 1970b. Further studies on marine turtles on Tonga
land, IV. Lammergeyuer 12: 26-36.
_________ . 1971a. The marine turtles of Tongaland, V. Lam
mergeyer 13: 7-24.
__________ 1971b. Sea turtle research and conservation in south
east Africa. A status report. Oceanographic Research Inst.,
Durban. 11 pp.
_________ . 1972. The marine turtles of Tongaland, VI. Lammer
geyer 15: 15-26.
_________ . 1974a. The sea turtles of south-east Africa. I. Status,
morphology and populations. Oceanographic Research In
stitute, Durban. Investigational Report, No. 35: 144 p.
_________ . 1974b. The sea turtles of south-east Africa. II. The
biology of the Tongaland loggerhead turtle Caretta caretta
L. with comments on the leatherback turtle Dermochelys
coriacea L. and the green turtle Chelonia mydas L. in the
study region. Oceanographic Research Institute, Durban.
Investigational Report No. 36 : 96 pp.
_________ . A.J. Bass and M.T. Mentis. 1967. Further studies on
marine turtles in Tongaland, 1. Lammergeyer 7, 5-54.
_________ . and M.T. Mentis. 1967. Further studies on marine
turtles in Tongaland, II. Lammergeyer 7, 55-72.
Kaufmann, R. 1966. Das Vorkommen von Meeresschildkroten
in Kolumbien and ihre Nutzung als Nahrungsquelle. Natur
und Museum, 96(2). Frankfurt a. M. 1.2 : 44-49.
__________ 1967. Wachstumsraten in Gefangenschaft gehalt-
ener Merresschildkroten. Mitt. Inst. Colombo-Aleman In
vest. Cient. 1: 65-72.
_________ . 1968. Zur Brutbiologie der Meeresschildkrote
Caretta caretta caretta L. Mitt. Inst. Colombo-Aleman In
vest. Cient. 2 : 45-56.
__________. 1972. Wachstumsraten in Gefangenschaft gehalt-
ener Meeresschildkroten. II. Mitt. Inst. Colombo-Aleman
Invest. Cient. 6: 105-112.
Klukas, R.W. 1967 ms. Factors affecting nesting success of log
gerhead turtles at Cape Sable, Everglades National Park.
Report on a phase of RSP-7, Everglades National Park.
57 pp.
LeBuff, C.R. 1969. The marine turtles of Sanibel and Captiva
Islands, Florida. Sanibel-Captiva Cons. Found., Spec. Pub.
No. 1. 13 pp.
746
__________and R.W. Beatty. 1971. Some aspects of nesting of the

loggerhead turtle, Caretta caretta caretta (Linne) on the
Gulf Coast of Florida. Herpetologica, 27(2): 153-156.
Medem, F. 1962. Estudio sobre tortugas marinas. Informe
sob re la comision realizada en la costa atlantica. Corpora-
cion Autonoma Regional de los Valles del Magdalena y del
Sinu. 12 pp.
Nishimura, S. 1967. The loggerhead turtles in Japan and neigh
boring waters (Testudinata: Cheloniidae). Publ. Seto
Marine Biol. Lab. xv(l): 19-35.
Parker, G.H. 1929. The growth of the loggerhead turtle. Amer.
Naturalist, 63: 367-373.
Rainey, W.E. and P.C.H. Pritchard. 1972. Distribution and
management of Caribbean sea turtles. Contribution no. 105,
Virgin Islands Ecological Research Station, Caribbean Re
search Institute, College of the Virgin Islands, St. Thomas.
10 pp.
Roth, C.E. 1973. Caretta. The Curious Naturalist, XII(7): 1-13.
Routa, R.A. 1967. Sea turtle nest survey of Hutchinson Island,
Florida. Quart. Jour. Fla. Acad. Sci. 30(4): 287-294.
Tufts, C.E. 1972. Report on the Buritaca Marine Turtle Nesting
Reserve with emphasis on biological data from “Operacion
Tortuga 1972” and Recommendations for the Future, ms.
73 pp.
__________ 1973. Operacion Tortuga. Informe Final del Proyec-
to 1973. 40 pp.
Uchida, I. 1967. On the growth of the loggerhead turtle, Caretta
caretta, under rearing conditions. Bull. Jap. Soc. Sci. Fish.
33(6): 497-507.
__________. 1973. Pacific loggerhead turtle—and its mystery of
oceanic life. Anima, No. 3: 5-17, 79.
747
www.Ebook777.com
Chapter 10
SIDE-NECKED TURTLES
The earliest ancestral turtles, which we now call amphiche-
lids, were unable to withdraw the head into the shell. However, as
time went by, natural selection corrected this deficiency independ
ently in two different amphichelid stocks. One of these stocks
evolved a mechanism which retracted the neck in a vertical plane
so that the head was still pointing forward even when fully retrac
ted, while in the other stock the head was pulled under the over
hanging front of the carapace by a lateral flexion, so that the
retracted head was pointed sideways. These two groups are now
known as the Cryptodira and the Pleurodira. The Cryptodira in
clude the vast majority of living turtles, while the Pleurodira con
sists of two families containing together about fifty species which
are the subject of the present chapter.
The Cryptodira have their area of greatest diversity and abun
dance in the northern hemisphere, while the Pleurodira are nearly
confined to the southern hemisphere. In the Cretaceous and early
Tertiary side-necked turtles were found in the northern continents
as well, but they were apparently displaced by the more progres
sive cryptodires in North America, Europe, and Asia. A disturbed
or attacked side-necked turtle can only protect itself by pushing its
snout further into its shoulder, and the whole side of the head and
neck remains exposed. This vulnerability has precluded the evolu
tion of terrestrial side-necks, since carnivorous mammals would
make short work of such slow-moving and poorly-protected prey,
and has also made side-neck turtles the losers in any direct compe
tition with cryptodires. Today, side-necks are found in Australia,
where they are almost the only non-marine turtles present, in
South America east of the Andes, in sub-Saharan Africa, and in
Madagascar. In the last three areas true tortoises (Testudinidae)
are present in some diversity but obviously provide no competi
tion. Emydid turtles are either very few (in South America) or un
represented (sub-Saharan Africa and Madagascar). In Africa,
several soft-shelled turtles are present, but soft-shells all over the
world appear to occupy an environmental niche which does not
preclude their occupying the same bodies of water as turtles of
other families. Another factor which may explain the survival of
748
side-necks in Africa is the evolution in all but one of the species of a

plastral hinge which enables the anterior shell opening to be closed
completely.
The sideways-bending neck of pleurodiran turtles reflects in
the cervical vertebrae, which have lateral processes for muscle
attachment and never show ginglymoidy or lateral doubling of the
articular surfaces between vertebrae. This doubling is found in
cryptodiran turtles, and its function is presumably to prevent the
neck from bending sideways when being withdrawn. In side-neck
turtles, vertical bending of the neck is to be avoided, and this is at
least minimized by a vertical elongation of the proximal cervical
articular surfaces. Other ways in which side-necks are distin
guished from cryptodires include the presence of an intergular
scute in all side-necks, but only in the sea turtles among crypto
dires, and in the sutural union of the pelvis with both the carapace
and the plastron. The pelvis in side-necks consists of two columns
braced between the shells, with a socket for the femur about half
way up, and with pubis and ischium bones on each side which
approach each other mesially but do not meet. The pelvis of a
cryptodiran turtle is a separate structure with no bony connection
with either carapace or plastron, but in which both ischia and
pubes do meet on the midline and become fused in adult speci
mens.
The two recognized families of pleurodires are the Pelomedu-
sidae and the Chelidae. The Pelomedusidae always have meso-
plastral bones and have the skull roof emarginated (if at all) from
behind. The Chelidae lack mesoplastra and have the skull roof
emarginated from below.
FAMILY PELOMEDUSIDAE
The family Pelomedusidae currently includes five genera:
Podocnemis, Peltocephalus, and Erymnochelys, which form the
subfamily Podocneminae, and Pelusios and Pelomedusa, which
form the Pelomedusinae. Pelusios is immediately recognizable by
the hinged plastron with mesoplastra meeting on the midline.
Pelomedusa differs from the others in having a very poorly
ossified plastron and five claws on the hind feet (only four in the
Podocneminae). Peltocephalus is distinguished from Podocnemis
by many characters, most of which reside in the skull. These
include the absence of a groove between the eyes, the low lateral
placement of the eyes, the hooked jaws, the completely
roofed-over temporal region, the simple ridging on the maxilla,
and many more technical features—as well as the reverse sexual
dimorphism (males bigger than females) and the unique
chromosome number (2n = 26). Erymnochelys is superficially
somewhat similar to Peltocephalus, but differs in having the same
749
www.Ebook777.com
chromosome number as Podocnemis as well as serological
similarities to that genus and also several unique characters,
including the pattern of cervical articulation, the presence of an os
centrale in the foot, and an iliac-carapacial connection that
impinges upon the suprapygal.
GENUS PODOCNEMIS
The genus Podocnemis derives its name from the enlarged,
greave-like scales on the hind feet. The skull in all the species is
extensively roofed over with bone, but some emargination from
both behind and below may be present. This is the most ancient
turtle genus extant, and at one time it was one of the most wide
spread. The genus is known from the late Cretaceous onward in
South America, and from the Eocene to the Pleistocene in Africa.
The genus prospers today in some diversity in South Amer
ica. The displacement of Podocnemis from Africa may well have
been the result of competition from the modern side-neck genus
Pelusios, which is not known from deposits earlier than the Mio
cene, but which today is widespread in sub-Saharan Africa, and
750
has reached Madagascar so recently that the form on that island is

indistinguishable from one of the mainland African populations.
Pelusios, having a good plastral hinge, is effectively as well pro
tected as any cryptodire, and this may account for the present suc
cess and recent expansion on the genus. Its arrival in Madagascar
may ultimately doom the endemic Erymnochelys species.
Podocnemis expansa is commonly known as the arrau river
turtle or charapa, or in Brazil the females are called iurara and
the male capitari. This is the largest of the pleurodires, mature
females of average size being two feet long and weighing about 70
pounds, while the largest individuals reach a carapace length of 35
inches and a weight of 200 pounds. Hatchlings, which are about two
inches in length, have uniform brown carapaces with very wide
central scutes—the middle three centrals are two to three times as
broad as they are long. The plastron, indented behind, is uniform
grey in color. The snout is pointed and a strong groove runs be
tween the rather close-set eyes down to the nostrils. The ground
color of the head is black with lemon yellow markings—a pair of
spots on the interparietal scale and another on each side of the
head between the eye and the tympanum. A narrow yellow line
outlines the upper margin of each orbit. As growth proceeds, the
central scutes get relatively narrower and the head pattern breaks
up; the first stage of the breaking-up process is marked by the
appearance of a black spot in the middle of each interparietal
blotch. The jaws are relatively pale in color in young specimens,
and a pair of small barbels is present just behind the symphysis of
the lower jaw.
Large female arrau turtles have very wide and distinctly flat
tened shells, sometimes with a shallow groove along the midline,
and in most individuals there are extensive areas of black on the
plastron. Males are much smaller and considerably less plentiful
than the females. The arrau has a tremendously wide distribution
embracing the entire Amazon system of Brazil, reaching Bolivia
via the Madeira River, and Colombia and Peru via the Solimoes,
or Upper Amazon. In Colombia it is found in the Putumayo and
Caqueta as well as the Solimoes. The species is found throughout
the Orinoco system, which is not unexpected since a branch of the
Upper Orinoco, the Casiquiare, flows into the Amazon. Individuals
are sometimes washed up in Trinidad by the Orinoco in flood. The
arrau also reaches the upper reaches of the Essequibo system in
Guyana; the two river systems do not come into actual confluence,
but in some places the tributaries are only a few hundred yards
apart. To the best of my knowledge the species is not found in Suri
nam. The arrau is entirely herbivorous when mature and shows a
number of interesting parallelisms with sea turtles. Large size is
one of these, and the habit of congregating in certain limited areas
751
www.Ebook777.com
to nest in another. Egg clutches are very large, complements of
over 100 eggs not being unusual. The turtles are obliged to con
gregate in great numbers for nesting, since nesting can only take
place in the dry season (September to November in Brazil, March
to April in Venezuela) when large sandbanks are exposed; and
since these sandbanks are relatively few in number, turtles con
gregate from many miles of river to nest on each one. Other
parallels with sea turtles are the spherical, soft-shelled eggs and
the habit of nesting under cover of darkness. The arrau has been
greatly reduced in numbers by excessive exploitation for a great
number of years. Even in the mid-nineteenth century, the
naturalist Henry Walter Bates estimated that 48 million eggs were
utilized from the Upper Amazon alone. The oil from the eggs is us
ed for lamps, for lubrication, for cooking, and for many other pur
poses. The adult turtles are killed in great numbers for food.
Legislation restricts exploitation of expansa in many parts of
Brazil, but enforcement is not universally effective. In Venezuela,
there is apparently some control over the number of turtles taken
on the four remaining Orinoco nesting grounds (Isla Pararuma,
Playa del Medio, Playa Blanca, and Cabullarito), but the numbers
taken may still be excessive. Transplantation experiments are be
ing undertaken, and new populations have apparently been in
troduced successfully into Lake Valencia in the Andean Highlands;
young specimens taken to the lake have grown up and nested on ar
tificial beaches. Tagging of wild turtles by the Venezuelans has
shown that a lapse of four years separates nesting seasons of in
dividual turtles.
Despite the breadth of its range, Podocnemis expansa is now
an officially endangered species; uncontrolled exploitation of the
females and eggs on the nesting beaches has virtually eliminated
the species from the entire Upper Amazon area of Brazil. The two
best surviving nesting areas are on the Orinoco in Venezuela near
the Colombian border and on the Rio Trombetas in Brazil south of
Guyana. Juhani Ojasti has documented the decline of the turtles
using the Orinoco Beach, from an estimated 330,000 in 1799 and
1800, to 123,600 in 1945, 36,100 in 1956, and only 13,800 in 1969. Some
of the most important beaches in Brazil and Venezuela are now
protected, but exploitation continues elsewhere.
Podocnemis unifilis, which is known as ‘teracay’ in Colombia,
is somewhat similar to P. expansa but does not reach such a large
size; the usual adult female is about fifteen inches long, though the
record length is 27 inches. The carapace is more domed than that
of expansa and does not develop a median groove. There is usually
only one barbel on the chin. The head markings of the juvenile are
bright orange-yellow in color. There is a median spot of this color
on the top of the snout, one on each side of the head in front of the
752
eye, one extending down the side of the head from the rear of the
eye to the lower jaw, one above the tympanum, and one on the
lateral edge of the interparietal scale. Other small spots may also
be present. The Interparietal scale is elongate, but the parietal
scales usually meet behind it. Suboculars are usually present,
separating the parietals from the maxillaries. The plastron is pale
yellowish-grey in hatchlings, and the soft parts are grey. Irregular
black blotches may be present on the plastron of the adult. As with
P. expansa, the central scutes are very wide at birth but become
relatively narrower with growth. Podocnemis unifilis occupies
Juvenile Podocnemis unifilis, an attactive side-neck once popular in the pet

trade. Photo by R.J. Church.
753
www.Ebook777.com
much the same gross range as P. expansa but appears to be more
of a lake dweller than the latter species, particularly when imma
ture. The reproductive patterns are very different, uni fills normal
ly laying only 15 to 25 eggs, exceptionally as many as 40. The eggs
are elongate, being about 1.6 inches by 1.1 inches in size, and more
than one clutch is produced in a season. The nesting season varies
greatly depending on the locality; in the Amazonas it is from July
to September, in the Llanos Orientales and the Guayabero-Guavi-
are River from late December to mid-February, in the upper and
middle Putumayo from November to January, in the lower Caque-
ta mainly in October, and in the upper Caqueta in November and
December. The sexes are readily distinguishable; females are
larger than the males, have short tails, and have light buff or
brown heads with black eyes. Males do not exceed a carapace
length of fourteen inches and more or less retain the yellow-spot
ted head and greenish eyes of the juveniles.
In captivity unifilis is less acutely cold-sensitive than expansa
and erythrocephala. This may reflect the somewhat wider temper
ature fluctuations that would be expected in the ponds and lakes
inhabited by unifilis than in the huge rivers inhabited by the other
two species.
Podocnemis erythrocephala, long known as Podocnemis cay-
ennensis, is found in certain ‘black water’ tributaries of the Ama
zon, principally the Rio Negro, but there are also a few isolated
records from the Rio Canuma (a small tributary of the Rio
Madeira, south of the Amazon), Tefe, and Santarem. Museum spe
cimens labeled Belem and Cayenne almost certainly were merely
shipped to France from those ports and did not originate there.
The most common vernacular name for this species in Brazil is
irapuca, while in English it is commonly called “red-headed Ama
zon side-neck.” Hatchlings are immediately recognizable by the
brilliant red or orange-red markings on the head—one between the
eyes extending down to the nostrils, another forming a triangular
band on the anteromedial part of the mandible, and a broad band
across the back of the head covering the greater part of the parie
tal and interparietal scales. This beautiful coloration is retained
throughout life by the males, but in adult females the head
becomes a dull brown. The ground color of the head in males and
young is dark brown. The neck and dorsal limb surfaces are usual
ly dark grey; the ventral surfaces vary from whitish-grey to dark
grey. The carapace is chestnut or rich brown in color, with a nar
row orange or red rim. The juvenile plastron may have an orange
or pinkish tint.
The carapace of erythrocephala is moderately domed and is
distinctly expanded posteriorly, especially in adults. The vertebral
keel is most prominent on the second and third vertebrals. A weak
754
nuchal indentation is usually present. The first marginal may be

longer than wide and is relatively more elongate than in other
members of the genus. The front plastral lobe is always longer
than the hind lobe, but the width of the two lobes is approximately
the same. The intergular scute is always longer than the gulars,
while the width of the intergular is almost always less than the
length of the intergular-gular seam. Adult males have a large
semicircular notch in the anal region of the plastron; this notch is
smaller and triangular In adult females. The head of erythroceph
ala is characterized by the grooved forehead and the very wide
interparietal scale, with the parietals meeting behind in almost all
cases. Subocular scales are almost invariably present, and the
masseteric scale does not reach the orbit. There are always two
barbels on the chin and usually three enlarged scales on the hind
foot. This is the smallest species of Podocnemis, the record shell
length being 12.6”. Hatchlings are about 1.6 inches in length.
Podocnemis erythrocephala nests from late August to early
November in the Rio Negro. Nesting usually takes place at night in
sandy, brush-covered areas; the eggs are elongate and either
hard-shelled or slightly flexible. The adults are primarily herbi
vorous, feeding on aquatic plants and fallen fruit; however, they
may also be caught on lines baited with fish. This species is hunted
by local people in the Rio Negro and is caught by harpooning, by
net, and on baited line. Females may also be caught on land when
they venture ashore to nest.
Podocnemis sextuberculata, known in Brazil as aiaca or putiu,
reaches a maximum length of only about one foot. This species is
known from the Amazon, the Putumayo, and the Caqueta River
systems. The carapace is dark in color and is distinctly convex, the
vertebral keel being raised into a swelling at the rear of the second
central. The plastron is straight-sided, indentate behind, and in
young specimens six prominent tubercles are present—one on
each pectoral, abdominal, and femoral scute. The entire underside
of the animal is grey in color. The head is grey, without markings
in adults, but with several small white spots in juveniles. There is a
deep groove between the eyes, but this groove is not continued be
low the nostrils. The jaws are broad, the parietal scutes are sepa
rated by the interparietal, and either one or two barbels may be
present on the chin. This is a carnivorous species and may feed on
fish in the wild state. Nesting is said to take place in July in the
upper Amazon, in November and December in the Putumayo, and
in October in the Caqueta. Recorded clutches numbered between 8
and 13 eggs; the nests are made both on the shores of lakes and on
sandy river beaches.
The other two South American Podocnemis species are both
circumscribed in range and rather poorly known. P. lewyana is
755
www.Ebook777.com
Podocnemis lewyana, an uncommon side-neck from Colombia. Both the
carapace and plastron are darkly colored. Specimen from the Lantz collec
tion.
Podocnemis sextuberculata from Leticia, Colombia. Notice the groove be

tween the eyes.
found in the Magdalena and Sinu Rivers in Colombia. It has a low

carapace of a very dark color and reaches a length of about sixteen
inches. The plastron is dark olive-brown. The head is rather nar
row, with a groove between the eyes, and there are two chin bar
bels. Subocular scutes are present, and the interparietal is broadly
heart-shaped, with the parietals meeting behind it. The skin of the
soft parts and head is grey, with pale yellowish-buff bands on each
side behind the eye. The interparietal scale is always uniformly
dark. This is the only member of the genus found north of the Cor
dillera Oriental. A peculiar feature, recently pointed out by Roger
Wood, is that there is no suprapygal bone in lewyana, the eighth
pleurals being triangular rather than trapezoidal.
Podocnemis vogli is an abundant species in the llanos of Vene
zuela as well as in the Departamento de Meta in Colombia, but in
spite of this it was first introduced to science as recently as 1935.
The local names for this species in Venezuela are galapago and
gurruna, while in Colombia it is known as sabanera. The carapace
is depressed, virtually keelless, and not expanded posteriorly. The
usual adult length is about twelve inches, but a shell in my collec
tion from the Apure River is fourteen and a quarter inches long.
The sexes differ in the shape and depth of the anal notch in the
plastron as well as in tail length and thickness and in size.
As in most of the species of the genus, Podocnemis vogli is somberly col

ored when adult. It nests away from rivers. Photo by Deuley.
757
www.Ebook777.com
The head is rather broad; the interparietal scale is elongate,
though the parietals meet behind it, and large suboculars are pres
ent. The maxillary scale is light posteriorly, dark in the middle
and anteriorly. The upper jaw is feebly notched, and the jaw sur
faces bear three roughened and denticulate ridges. A large vomer
is present, extending from the premaxillae to the palatines. Two
chin barbels are present and there are three enlarged scales on
each hind foot.
The carapace of vogli is brown with indistinct yellow spots;
the plastron has a yellow ground color with black quadrangular
blotches on each scute. Young specimens have five or seven light
yellow spots on the head, but these fade with age, especially in
females. This species is found both in moving waters and in lakes
and may even be found in temporary pools. They frequently
emerge from the water to bask in the sun on emergent logs and
snags. This is a primarily vegetarian species, but captive speci
mens feed happily upon fish and meat. Nesting takes place from
late October to early January and appears to be under some lunar
influence. Instead of nesting in sandbanks and river beaches as is
typical of some other members of this genus, vogli females may
wander distances of many hundreds of yards on to the savannas
for nesting. The excavation of the nest cavity is assisted by the
excretion of cloacal water. Recorded clutches numbered between
7 and 17 eggs, and there is evidence that there may be two nestings
during a season. On the Orinoco coast, however, in Estado Guar-
ico, as many as twenty eggs may be laid, and the season extends
into February. The eggs measure between 3.7 and 4.8 cm in length
and between 2.1 and 2.8 cm in width. The hatchlings appear in late
April and May. The eggs and young are preyed upon by caracaras
and tegus, while adults are subject to predation by jaguars.
GENUS PELTOCEPHALUS
Peltocephalus tracaxa is a distinctive, relatively little-known

species which for many years was called Podocnemis dumeriliana.
However, many characters demonstrate that it is generically
distinct from Podocnemis. The name dumeriliana cannot be used
as the type specimen in the Paris Museum proves to be a
Podocnemis unifilis. *
The carapace is convex, with a vertebral keel that is most
marked on the last two central scutes and reaches a maximum
length of about nineteen inches. The carapace is dark greyish
brown in color, the marginals edged with light yellowish. The
plastron is dull yellowish brown, clouded with grey, and with a light
yellowish periphery. The intergular is larger than each gular, the
interhumeral and interpectoral seams are of comparable length,
*J. Fretey, 1977. Thesis, University of Paris.

The generic characters of Peltocephalus tracaxa are mostly internal, but the
very large head lacking a groove between the eyes is fairly distinctive.
and the interfemoral seam is the longest of the plastron. The head
is powerfully constructed with strongly hooked jaws and shows
similarities to that of Macroclemys or Platysternon. There is no
groove between the eyes. The skull roof is completely roofed-over
as is that of a sea turtle. The parietal scales are well separated by
the interparietal, and on each side the parietal is separated from
the maxillary scale by the masseteric. In color the head Is dar
greyish brown, without bright markings, though there are vague
yellowish mottlings between the eye and the tympanum, and a
blackish spot below the eye Is flanked by more yellowish mottlings.
A single chin barbel Is present. The soft parts are greyish.
The overall distribution of tracaxa is still incompletely
understood. It only reaches high population densities in certain
black-water rivers, such as the Rio Negro and the Atapabo, on the
Colombia-Venezuela border. Medem reports that it is found in the
Caqueta and Putuihayo River systems in Amazonian Colombia, as
well as the Uper Rio Tomo, the Isana, the Guainia, and the Inirida.
The species is not known from either Guyana or Surinam, but has
759
www.Ebook777.com
been reported from the Savane Ouassa, near the border between
Brazil and French Guiana. It is probably widespread, but spottily
distributed, in the Brazilian Amazon, having been reported from
points as far apart as the Rio Trombetas and Codajas (Upper
Amazon), and it apparently extends into Amazonian Peru. In
Central Venezuela, north of the Atabapo, it is rare but is known
from the Sinaruco, Potrerito, and Capanaparo Rivers.
Peltocephalus tracaxa is usually called cabezon, or cabecudo,
both names referring to the large head. The formidable hooked
jaws give the impression that this species must be a powerful
predator, but the scant evidence on hand suggests that this is not
the case; those stomachs that have been examined contained
mostly fruits of various species of palm. Nesting is said to take
place in July and August in the Upper Amazon. The eggs are very
large—about 53 to 57 mm in length and 35 to 38 mm in width—and
relatively few in number; usually nests contain only about seven to
twelve eggs, though up to 25 may be deposited on occasion. Instead
of nesting in sand banks, as do other species of the Podocneminae,
tracaxa wanders off into flooded forest areas and nests in any
piece of ground that is sufficiently above water level. Eggs and
hatchlings are therefore very hard to find.
GENUS ERYMNOCHELYS
Erymnochelys madagascariensis was long considered to be a

member of the genus Podocnemis and is the only living member of
the Podocneminae to inhabit the Old World. It shows a number of
similarities to Peltocephalus, but for reasons outlined earlier in the
chapter the species is distinct enough to warrant generic recognition.
Reaching to about seventeen inches, the carapace is oval,
moderately depressed, and keeled in the young; the carapace
margins are expanded and reverted. Radiating striae are evident on
the carapace scutes sis well as fine black dots and striations. The
plastron is large but distinctly narrower posteriorly than anteriorly.
The intergular scute is very small, triangular In shape, and allows the
gulars to meet behind it (in the South American relatives the in
tergular completely separates the gulars and partially separates the
humerals). The ground color of the plastron is yellow, but brown
pigmentation is often present as well. The upper jaw is slightly hooked
and there is no groove between the eyes. The interparietal scale
tapers posteriorly, but the parietals meet behind it. The masseteric
scale usually reaches the maxillary scale, but sometimes suboculars
intervene. There is usually only a single chin barbel, but sometimes
there are two, and there are three enlarged scales on the border of
each foot.
760
GENUS PELUSIOS
Turtles of the genus Pelusios are found over much of sub-
Saharan Africa. The genus is immediately recognized by the hinge
across the plastron which allows the front portion to be raised for
protection of the head and forelimbs. The hind part of the plastron
is completely rigid; not only is the bridge well-developed, but the
pelvis provides a fixed connection between the hind parts of the
carapace and the plastron. This is the only living turtle genus in
which mesially contiguous mesoplastral bones are present in the
plastron. As in Podocnemis and Pelomedusa, the hindmost pair of
pleural bones has a median line of contact. The head is blunter
than that of Podocnemis, and the temporal region of the skull is
deeply emarginated from behind. The superficial appearance of
the skull is somewhat similar to that of many emydid turtles,
though close examination reveals many subtle differences. As in
Podocnemis, the head is covered with a small number of very
large, symmetrical shields.
The number of valid species of Pelusios is debatable; some
authorities only admit the existence of five species, others as
many as ten. In this book I have followed the ‘splitters’ rather than
the ‘lumpers,’ mainly because the work of some modem ‘split
ters,’ such as Raymond Laurent, has been confirmed by the serio-
logical studies of Wayne Frair and others. But it should be men
tioned that, while the species gabonensis, adansoni, nanus, sinua-
tus, and subniger are definitely valid, the other species listed be
low are looked upon with various degrees of dubiousness by some
competent authorities.
The most distinctively marked member of this genus is the
Gaboon turtle, Pelusios gabonensis. This species is found in tropi
cal West Africa, having been recorded from Guinea, Liberia,
Ghana, Nigeria, Cameroon, Gabon, Zaire, and Cabinda. The cara
pace is depressed and light yellow-brown in young specimens, with
a prominent slightly irregular black vertebral line. The plastron is
large and typically almost entirely black except for light edges to
each scute, but in some specimens buff or yellow areas may be
more extensive. The plastral hinge is set well back so that nearly
half of the plastron is moveable; the anterior lobe is thus more
than twice the length of the interabdominal seam. The head is
broad and flat, light brown above with a black median streak that
broadens on top of the head to reach both orbits. The chin, throat,
and soft skin are light in color, but the feet are greyish. Large in
dividuals are not as clearly marked as juveniles and have power
ful jaws, the upper jaw being mesially hooked. A pair of barbels is
present on the chin. The scales on the front face of the forelimbs
are irregular in size. The maximum length is about one foot. The
Gaboon turtle inhabits swampy places and small streams when
761
www.Ebook777.com
young, but moves into larger rivers when mature. It is carnivorous
and insectivorous in captivity. George Cansdale reports that the
number of eggs laid by this species is ‘probably about a dozen.’
The serrated turtle, Pelusios sinuatus, is found in East Africa
from Somalia south through Kenya, Tanzania, Rwanda, Burundi,
Mocambique, Malawi, Zambia, Rhodesia, and Botswana to Zulu-
land, Transvaal, and Natal. This is the largest member of the
genus, reaching a maximum length of over fifteen inches and a
weight of perhaps twenty pounds. The carapace is black and mod
erately depressed in young and half-grown individuals, but quite
deep in some adults. The hind margin is strongly serrated or wavy
in all except the oldest individuals. Each of the central scutes is
slightly swollen toward its hind margin. The front lobe of the plas
tron is relatively short—shorter than the interabdominal seam.
The underside of the marginals, the bridges, and the periphery of
the plastron are black, but the central part of the plastron bears a
characteristic symmetrical unpigmented area with a strongly
angular outline, a wedge-shaped section of this unpigmented area
extending close to the outer edge of each of the pectorals, abdomi
nals, and femorals. The dorsal parts of the head are yellowish,
orange-yellow, or pale olive, finely speckled with black or dark
brown. The iris of the eye is light yellowish in color. The soft parts
are pale buff-grey, becoming grey on the feet. A pair of barbels is
present on the chin. All the toes are webbed. This species is carniv
orous, eating insects, snails, and fish. It has been recorded from
Pelusios sinuatus, the serrated turtle, showing the deeply wavy posterior
margin of the carapace.
lakes and large rivers in the coastal belt and upland savanna up to
an altitude of 5,000 feet.
Adanson’s turtle, Pelusios adansoni, is found across northern
tropical Africa, from the White Nile in the Sudan through Chad,
Cameroon, Nigeria, Mali, Liberia, Senegal, and the Gambia to the
Cape Verde Islands. This is a small species, large adults being
only about seven inches long. This species shares with gabonensis
Adanson’s turtle,
Pelusios adan
soni,of northern
Tropical Africa has
a nearly unmarked
plastron. Speci
men from the
Lantz collection.
the long anterior lobe of the plastron; the plastron in front of the
hinge is more than twice the length of the interabdominal seam.
The carapace is somewhat depressed, wider posteriorly than an
teriorly, rounded behind, and with a keel on the first four central
scutes even in adults. The upper surface of the head is ver-
miculated with light and dark markings. The carapace is dark, but
the plastron is predominantly or completely unpigmented. The
head is broad, the snout short, and the upper jaw neither hooked
nor bicuspid. Two barbels are present on the chin. A specimen col
lected in March in the White Nile contained seven eggs, each about
1.2x0.8 inches in size. The habitat is said to embrace rivers outside
the rainforest.
763
www.Ebook777.com
The western African Pelusios niger is heavily vermiculated with black on the
head and has a uniform blackish plastron.
Pelusios nanus, described in 1956, appears to be a southern rel

ative of adansoni; it has been recorded from Angola and Zaire.
This is the smallest species of the genus, being adult at less than
five inches. It differs from adansoni in the narrower third central
scute; however, exact ratios cannot be given as they alter with
age, and several other ratios that formerly were supposed to dif
ferentiate the species have now been shown to be too variable to
use.
Pelusios niger is another West African species, known defi
nitely from Ivory Coast, Nigeria, Gabon, and Cameroon, and pos
sibly extending from Senegal and the Gambia to Angola. Both car
apace and plastron are black, but the seams are often white. The
head is narrower than in many other species of Pelusios, and the
snout is somewhat pointed. The jaws and upper parts of the head
are yellowish-grey, with strong black vermiculations. The soft
parts are pale, often brilliant white near the periphery of the plas
tron. The species is distinguished from all subspecies of P. casta-
neus by the wider intergular border and narrower femoral border
(ratio of these two quantities about 0.425:1 in niger, about 0.28:1 in
castaneus). A median keel is present on the carapace, particularly
elevated at the rear of the third and fourth centrals. The anterior
lobe of the plastron is only slightly longer than the interabdominal
seam, and, unlike the condition in subniger, the plastron is not
markedly constricted just behind the bridge. Adults reach a cara
pace length of eight or nine inches.
Several of the species considered as full species in the present
discussion are commonly synonymized with Pelusios subniger.
However, P. subniger in the strict sense is an East African form,
known from Ethiopia south to Transvaal, and also recorded from
Madagascar, Mauritius, and the Seychelles. The island popula
tions are not distinguishable from those on the mainland, and it is
not inconceivable that they reached the islands through human
agency, though there is not actual evidence for this. The carapace
is unkeeled, with a smooth hind margin and dark grey-brown in
color. The plastron may be dark brown or black, entirely yellow,
or yellow with some more or less symmetrical dark markings. The
hind lobe of the plastron is strongly constricted just behind the
bridge. The front lobe of the plastron is only a little longer than the
interabdominal seam. The intergular scute is moderately broad
but narrows anteriorly, and a strong, angular notch is present at
the rear of the plastron. The vent in male specimens is situated al
most at the tip of the tail. The head and limbs are a uniform grey
color without any vermiculated pattern on top of the head. On
casual inspection the eye appears uniformly dark, with no light
ring around the pupil.
765
www.Ebook777.com
Plastral view of
Pelusios subniger
of eastern Africa.
The color is very
variable.
Pelusios carinatus, from Zaire, is frequently confused with P.

subniger but differs in having a widening rather than a constric
tion of the plastron behind the bridge, a vermiculated or marbled
head pattern, and a yellow plastron with a black border on the
front lobe only. The carapace is strongly keeled and has a serrate
hind margin. Only a shallow indentation is present between the
femoral and anal scutes. The intergular is usually 75% or less of
the length of the combined interhumeral and interpectoral seams,
at least in adult specimens.
Pelusios castaneus is a widespread species known from the
Seychelles, Madagascar, Zambia, Pemba Island, Malawi,
Rwanda, Burundi, Angola, Zaire, Liberia, Sierra Leone, Portu
guese Guinea, and the Gambia. The species differs from subniger
in having a variegated head pattern, no or little plastral constric
tion behind the bridge, a narrower head, large sickle-shaped
scales on the forelimbs, a shorter intergular scute (usually about
766
equal in length to the combined interhumeral and interpectoral

seams in adults), and white-bordered irides in the eyes. Four sub
species have been described. The typical form, from coastal East
Africa, Madagascar, and Pemba Island, as well as the Seychelles,
is rather light in color, with a brown, yellowish brown, or chestnut
brown carapace and often a yellow plastron. Three of the races are
not known to exceed ten inches in carapace length, but chapini,
which is isolated from all the other races in northern Zaire,
reaches about fifteen inches and has relatively wider intergular,
gular, and abdominal scute borders, while in adult specimens the
femoral border is usually shorter than the abdominal border. Pelu
sios castaneus rhodesianus, from southern Zaire, Rwanda, Burun
di, Angola, Rhodesia, and Malawi, tends to be dark in color, often
with a completely black plastron. P.c. derbianus, from West Afri
ca (Liberia, Sierra Leone, Portuguese Guinea, and the Gambia,
but not Ghana and Nigeria, where niger is present), may be
weakly differentiated by the short interpectoral seam. A popula
tion of one or other of the subspecies of Pelusios castaneus is pres
ent on the islands of Guadeloupe and Marie Galante, in the French
West Indies. Although it is quite likely that these turtles were intro
duced to the islands by man, the interesting possibility remains
that natural colonization took place by way of the North Equa-
torian Current. Certainly Pelusios seems to show a greater ability
to colonize islands than most freshwater turtles.
www.Ebook777.com
Pelusios williamsi is a newly described species known from
the Upper Nile Basin. It is defined as ‘a large species with a black
and yellow plastron whose anterior lobe is of moderate length
(longer than the abdominal suture but shorter than twice this
length), differing from P. castaneus by its larger head, its larger
intergular, narrower third central, and shorter femoral border’
(Laurent). Two subspecies have been described: P.w. williamsi
from the lake Victoria region, and P.w. lutescens from Lake Al
bert, the Semliki River, Lake Edward, and their affluents. The
typical form has a predominantly black plastron (yellow with
some greyish or brownish spots in lutescens) and a relatively
longer intergular scute than lutescens.
Pelusios bechuanicus was originally described from a four-
inch juvenile from the Thamalakane River at Maun, Ngamiland,
Botswana. Loveridge considered the species inseparable from
subniger, but Laurent, with further material of the form from
Angola, has reinstated it. This species resembles many subniger in
its uniform dark brown plastron and posteriorly rounded cara
pace, but is similar to sinuatus in the protuberances on the central
scutes. The central scutes are reported to be longer than broad,
even in the young.
Specimen of Pelusios williamsi, rather recently described from the Nile

drainage. This is P. w. williamsi from the Lake Victoria region. Specimen from
768
Pelusio williamsi lutescens is from the Lake Edward - Lake Albert area.
Plastral view of Pelusios williamsi williamsi (left) and P. williamsi lutescens

(right) illustrating the color difference.
769
www.Ebook777.com
GENUS PELOMEDUSA
The helmeted turtle, Pelomedusa subrufa, is placed in a genus
apart from Pelusios because the plastron lacks a hinge and the
mesoplastra are small and widely separated. The species is found
over much of Africa, having been reported from the Sudan, Ethio
pia, Togoland, Ghana, Senegal, Mali, Nigeria, Cameroon, Eritrea,
Somalia, Uganda, Kenya, Tanzania, Mocambique, Malawi,
Rhodesia, Zambia, Botswana, Angola, Zaire, Transvaal, Orange
Free State, Cape Province, Natal, and many localities in Mada
gascar. Geographic variation within the species has not been re
corded, with the exception of a tendency for specimens from some
of the more northern localities to have separated pectoral scutes.
This separation appears to be universal in Eritrea, and such speci
mens have been designated as a distinct subspecies (olivacea).
The helmeted turtle is adult at a carapace length of six or seven
inches, but a male from Kingwilliamstown District, South Africa,
is reported to have had a shell approximately thirteen inches in
length. The carapace is depressed, some shade of brown in color,
and smooth or very slightly serrate behind. The plastron is yellow
ish or horn color, frequently with dark seams and various pigmen
ted areas on the scutes. The bridge is made up of about equal con
tributions of pectoral and abdominal scutes; in Pelusios the ab
dominals make up almost all of the bridge. The shell is very thin,
Pelomedusa subrufa, the helmeted turtle, is found over much of Africa. The
shell is flat and thin.
770
and a large fenestra in the bony plastron persists until adulthood.

The head and limbs are dark greyish or olivaceous above, pale yel
low or white below. This is a semi-aquatic species found in shallow
and temporary bodies of water or in marshes. When these dry up,
it estivates. It is known to bask on warm days in South Africa, but
appears not to do so in tropical Africa. It has been found up to an
altitude of 6,000 feet. This species appears to be unusually prolific
for a small turtle; a specimen which weighed about five pounds
was kept by W.H. Archer and laid 31 eggs on December 9,1963, and
42 about a year later, on December 22, 1964. Other recorded clut
ches of this species numbered 16 and 14 eggs. The eggs measure
about 1.5 inches by 1 inch and take a little more than three months
to hatch.
The helmeted turtle appears to be almost entirely carnivor
ous. It is not immediately obvious why a side-necked turtle without
a protective moveable plastral lobe has managed to survive in
great abundance in Africa today, while Podocnemis has become
extinct on that continent. Possibly the answer is connected with
the habitat versatility of the species and its ability to make over
land treks to exploit temporary aquatic situations.
FAMILY CHELIDAE
The other family of side-neck turtles, the Chelidae, shows a
number of advancements over the more primitive Pelomedusidae.
For example, the mesoplastra have been lost and the bony shell
mosaic of more primitive chelids (Chelus and Hydromedusa) is,
fortuitously, virtually identical to that of modem cryptodires.
Other chelid genera go even further in simplifying the arrange
ment of shell bones, reducing the number of neural bones to six or
less (Phrynops), or to zero (Platemys and most Australasian spe
cies). The skull is also more specialized than in the Pelomedusi
dae, the different genera showing a progressive emargination of
the temporal roof from below (not from behind, as in cryptodires).
In Pseudemydura the skull roof is almost intact, with only a slight
emargination. In Emydura, Elseya, Chelus, and Platemys exten
sive emargination is present. In Hydromedusa and Phrynops only
a narrow curved bar at the extreme rear of the skull remains, and
in Chelodina the temporal bar disappears completely, giving a
condition similar to that of certain emydids (e.g. Heosemys)
except that the condition was preceded evolutionarily by one in
which the temporal bar was at the rear of the skull instead of
behind the eye.
771
www.Ebook777.com
A prominent feature of chelids is the neck; several forms have
extremely long necks, and even in the shorter-necked forms the
lateral sigmoid flexure of the neck is much more conspicuous than
in the Pelomedusidae. The cervical vertebrae show a different
arrangement of articulation in the two families; in the Pelomedu
sidae the second cervical is biconvex (i.e. has convex articular
surfaces at both front and rear), while the third to eighth cervicals
are all procoelous (i.e. have a concavity in front and a convexity
behind). On the other hand, in the Chelidae the second to fourth
cervicals are opisthocoelous (i.e. with a convexity in front and a
concavity behind), the fifth is biconvex, the sixth is procoelous, the
seventh is biconcave, and the eighth biconvex.
At the present time the Chelidae are restricted to South Ameri
ca east of the Andes and to Australia and New Guinea. According
to Romer, the family was found in Europe and North Africa in the
Cretaceous, but Ernest Williams considers these fossils doubtful at
best, and Darlington concludes that there is no sure evidence that
chelids ever had a wider range than they do now.
GENUS CHELUS
Chelus fimbriatus, the famous matamata turtle, is one of the
most bizarre of all chelonians. It is a rather large species, reach-
772
ing a carapace length of about sixteen inches, found in many parts

of the Amazon system, in the Orinoco, and in several of the north
draining rivers in Guyana and French Guiana, though there ap
pears to be no record of the species from Surinam. It has been
recorded from Trinidad, this species and Phrynops (Mesoclem-
mys) gibba being the only chelids on that island.
The carapace of the matamata is exceedingly rough and has
three prominent tuberculate keels, the highest tubercles being
toward the rear of the median keel. Seven or eight neural bones
are present, and only the eighth pair of pleurals sometimes has a
median line of contact. The bridges are rather narrow, as is the en
tire plastron, and the rear plastral margin is deeply notched. The
head is so flat and distorted that a person seeing a matamata for
the first time usually has difficulty locating the eyes, which are ex
tremely small and situated very close to the snout. The snout itself
is extended forward into a long, very thin, snorkel-like breathing
tube. The head becomes much wider posteriorly, and the tympanic
membranes are exceedingly large and prominent. The neck is
long, very thick, and extremely muscular; the limbs, however, are
rather small and weak.
The matamata, Chelus fimbriata. Head view showing the long snout and
fringes. Photo by Ken Lucas, Steinhart Aquarium.
773
www.Ebook777.com
Adult matamatas are uniformly dark brown or black in color,
but young specimens are more colorful, with a rich brown cara
pace and plastron, and dark centers to the plastral scutes. Also, in
young specimens the underside of the neck is often marked with
three broad scarlet longitudinal bands separated by two black
bands. The mouth of the matamata is almost absurdly wide, but
the jaws are rather weak and the two halves of the lower jaw are
not fused in the middle. The two pairs of hyoid bones are fully as
large as the lower jaw bones, which they closely resemble in shape
—recalling the serial homology that actually exists between these
elements. The chin and sides of the head and neck of the matamata
are decorated with numerous branched filamentous appendages.
The strange specializations of the matamata are all in keeping
with its sedentary but predatory way of life. It occupies an ecologi
cal niche not dissimilar to that of the alligator snapping turtle
(Macroclemys) of North America. The matamata lies on the bot
tom of the water, moving so little that a thick growth of algae
forms on its rough shell, providing near-perfect camouflage. When
small fish swim near the head, they are sucked right into the ani
mal's gullet by a sudden opening of the mouth and expansion of the
huge neck—an expansion assisted by the well-developed hyoid
bones and the massive neck musculature. The turtle then closes its
mouth to a slit, allows the water to escape, and swallows the fish.
Some recent experiments by Peter Hartline demonstrate that
the chin and neck flaps of a matamata are highly innervated and
sensitive to slight water movements and pressure waves. This sen
sitivity apparently enables the matamata to locate fish swimming
near it in murky black-water rivers. Whether they also serve to
attract fish to the vicinity, in the same way as the vermiform lure
on the tongue of the alligator snapping turtle attracts fish, is not
known.
The eggs are spherical, have hard, brittle shells and measure
about 1.5 inches in diameter. Nesting takes place in October in
Amazonas, Colombia, and in November and December in the Put
umayo. Between 12 and 28 eggs may be laid at a time. A clutch in
cubated in Germany by Peter Hausmann took 208 days to hatch, on
average. The young have light tan-colored carapaces with a black
median line extending to the back of the head, a narrow V-shaped
marking on the top of the head pointing toward the snout, three
large black spots on each side of the carapace, and black curved
markings on the outer border of each of the marginal scutes.
No subspecies of Chelus fimbriatus have been described, but
Medem reports that specimens from different areas of Colombia
differ in coloration, especially on the underside of the neck.
Schmidt (1966) observed differences between individual mata
matas from Brazil, Peru and Colombia, but his conclusions were
774
suggestive rather than definitive, and a systematic approach is

essential before subspecies can be named.
GENUS HYDROMEDUSA
The genus Hydromedusa, while superficially very different
from Chelus, and similar in some features of external morphology
to Chelodina, is perhaps more closely related to the matamata
than to any other species. The genus is characterized by the
extremely long neck, somewhat flattened skull, presence of a
more-or-less complete series of neural bones (six to nine in num
ber), and the peculiar nuchal scute, which is unusually wide and is
situated completely behind the anterior marginal scutes, giving
the impression of an extra vertebral scute. The intergular scute is
large and completely separates the gulars; it also partially
separates the humerals.
Two species are recognized: Hydromedusa tectifera in
Argentina (from Rio de la Plata to Chaco and Formosa), Para
guay, Uruguay, and southern Brazil (states of Rio Grande do Sul
and Sao Paulo, and probably also Parana and Santa Catarina);
and H. maximiliani, which has a much smaller range, and judging
by the relative numbers in museum collections, appears to be a
much rarer animal—almost all known spe'cimens are from the
state of Sao Paulo, including the island of Sao Sebastiao, but the
species has also been reported from Parate, Rio de Janeiro. The
two species are easily distinguished. H. tectifera is a much larger
animal than H. maximiliani; the former may reach a carapace
length of 30 cm, while the latter is not known to exceed 21 cm. The
latter is also distinguished by the strong muscular fold that bulges
out on each side at the corner of the mouth ; no trace of this is pres
ent in tectifera. Roger Wood and R.T. J. Moody have also recently
described a feature of maximiliani that is completely unique
among turtles; the nuchal bone, like the nuchal scute, is inset from
the front margin of the carapace, so that the front pair of peri
pheral bones meets in front of it. In tectifera, as in other turtles,
the nuchal bone forms part of the front margin of the carapace.
Other features mentioned by Wood and Moody include the follow
ing: in tectifera there are usually six neural bones; pleurals 7-8
meet on the midline; there are 22 peripherals; the entoplastron
has a small rugose attachment area; the axillary notch is not sig
nificantly wider than the inguinal notch; the first pair of marginals
is much broader than long; the nuchal scute is as broad as the
combined width of the first pair of marginals; the last pair of mar
ginals is longer than broad ; and the shortest scutes along the mid
line of the plastron are the humerals or pectorals. In maximiliani
there are seven to nine neural bones; pleurals eight may meet in
the midline; there are 22-24 peripheral bones; the entoplastron has
775
www.Ebook777.com
Plastral view of Hydromedusa maximiliani showing the unmarked shell and
soft parts. Specimen from the Senckenberg Museum.
The extremely long neck of Hydromedusa tectifera is obvious in this photo of

a specimen from the Brownsville Zoo, Texas.
776
a relatively large rugose attachment area; the axillary notch is

considerably wider than the inguinal notch; the first pair of mar
ginals is almost as broad as long; the nuchal scute is broader than
the combined width of the first pair of marginals; the last pair of
marginals is broader than long; and the shortest scutes along the
midline of the plastron are the abdominals. Other differences in
clude the tendency of tectifera to retain an unossified fenestra in
the middle of the plastron into adulthood; the plastron of maximili
ani becomes fully ossified with maturity. Tectifera also tends to
have a deeper notch at the rear of the plastron than maximiliani.
There is some difference in coloration between the two spe
cies. The appearance of tectifera is typified by a young live speci
men from Uruguay in my possession, the carapace of which is
approximately five inches in length. The carapace is flat, very
dark brown—almost black—in color, with considerable surface
sculpturing. The shell does not appear to desquamate or flake in
this form, and growth annuli and radiating striae remain in evi
dence until they wear down. Carapaclal tubercles are well devel
oped on the vertebral and costal scutes in the very young; in the
five-inch specimen these are only really evident on the posterior
scutes—the last two vertebrals and the last costals—though the
areolae of all of the larger scutes are slightly tubercular. The dor
sal surfaces of the limbs, the head, and the exceedingly long neck
are a uniform dark grey though the tips of some of the tubercles on
the sides of the upper surface of the neck are yellow-buff. The plas
tron is almost uniform yellow-brown with distinct growth annuli on
each scute. The bridges and parts of the under-marginals are dark
brown. The lower edge of the dark pigmentation on the head is
marked by a black line that begins at the nostril and passes
through the eye and the tympanum before fading out on the neck.
Below this is a very conspicuous yellowish-white streak that com
mences on the upper mandible and extends the length of the neck.
The underside of the neck and throat are marked with complex
vermiculations of pale yellow and dark grey. The lower surfaces of
the feet are pale yellow with dark grey markings.
The appearance of maximiliani differs significantly from that
of tectifera, quite apart from the features mentioned earlier. The
shell is smoother and virtually lacks the median keel. The plastron
is brown in the adult, yellow spotted with brown in the young. The
upper surface of the head is covered with finely wrinkled skin, and
the lower surfaces of the head and neck are uniformly pale, the
pale area extending up to the upper mandible and the lower edge of
the tympanum.
The two species of South American snake-necked turtles,
which may later be thought distinct at the generic level, are
rarely imported into the United States but are quite plentiful in the
777
www.Ebook777.com
wild. H. maximiliani, with the much more restricted range of the
two, is often considered rare, but it is now known to exist in abund
ance on the island of Sao Sebastiao off the coast of the state of Sao
Paulo. The habits of the two species have found few observers.
Paul Mueller, who caught numbers of maximiliani on Ilha Sao
Sebastiao, found insect larvae and tadpoles of three species (Lep-
todactylus ocellatus, L. pentadactylus, and Bufo crucifer) in the
same impoundment and postulated that these might be food items
for the turtles. Freiberg writes that tectifera in Argentina eats
small fish and amphibians and in captivity will engulf pieces of
chopped fish with a rapid sucking action similar to that of the
matamata turtle. However, the preferred food is snails; its weak
jaws are ill adapted to crushing the shells of these animals, but it
methodically removes the operculum and cleans out the contents
by placing its snout and jaws right in the opening.
H. tectifera in the wild is reported by Freiberg to be very
nervous and shy and to retain these characteristics in captivity,
although the species is not hard to maintain in good health in a cap
tive situation. During the cold months tectifera hibernates at the
bottom of the water. The eggs measure about 34 x 20 mm. The
young present a bizarre appearance with their exceedingly rough,
tubercular shells. The sexes are rather similar, but the females
reach a larger size than the males, and the latter show a plastral
concavity centering on the midline of the femoral scutes.
A third member of the genus, Hydromedusa depressa, has
been described but is clearly a synonym of maximiliani; the shell
of the type specimen had been physically crushed at an early stage
of ontogeny and, although healed when the animal was collected,
showed numerous resultant deformities.
GENUS PLATEMYS
The genus Platemys includes three small South American spe
cies characterized by the complete absence of neural bones and
the presence of a median groove in the carapace flanked by two
blunt ridges. The most familiar species is the northern representa
tive, Platemys platycephala, commonly known as the flat-headed
flat-shelled turtle, or (in Guyana) as the twist-neck turtle or cashi-
pan. It has been recorded from all three Guianas, much of Amazo
nian and northern Brazil, Venezuela, Colombia, and eastern Ecua
dor and Peru. Platemys platycephala is adult at a length of about
six inches. The carapace is very flat, oval in outline, and has a
median groove which produces a strong convexity in the second,
third, and fourth centrals. The bridge is unossified, and the plas
tron is large and flat; the intergular scute is very wide, completely
separating the gulars and partially separating the humerals. The
limbs are rather weak and are covered with large smooth scales.
778
Platemys platycephala, the most familiar species of the genus, is recognized

by the shape of the carapace and the bright orange head.
The head is very flattened, and the upper part of the neck is
covered with pointed tubercles. The skin on the top of the head is
undivided. The snout is pointed, and there is a pair of small barbels
on the chin. In color the carapace is rich chestnut-brown with
black streaks covering part of the first costals and most of the sec
ond and third costals. The plastron is dark brown or black except
for a narrow yellow rim. The bridge is yellow with a broad longitu
dinal black streak, and the underside of the marginals is yellow,
sometimes with black markings. The top of the head is rather
bright orange, and this passes into yellow on the tubercular area
on top of the neck; otherwise the extremities are uniformly black
except for a couple of pointed yellow scales of the underside of the
ankles.
Platemys platycephala is basically aquatic but is quite often
found wandering on land after rain. This is the most frequently en
countered side-neck turtle in northern Guyana and Surinam and in
places reaches within a few miles of the sea. Breeding is not seaso
nal, and four to six eggs are laid at a time, often a long distance
from water. In captivity this species does well and lives for many
years, but its movements, muscle tone, and whole demeanor may
appear so feeble that it often appears to be on the point of expiring.
Platemys pallidipectoris is a slightly larger species than
platycephala, reaching a length of about eight inches. It was dis
covered in the Chaco of Argentina in 1946. It may be identified
779
www.Ebook777.com
Plastral view of the uncommon Argentine Platemys pallidipectoris. Photo by
Dr. Robert Mertens.
easily by the cluster of enlarged tubercles on the thigh between the

hind legs and the tail. The limbs are covered with enlarged scales
and the neck with coarse rounded tubercles. The plastron, al
though mainly black, has unpigmented areas on the outer part of
each scute.
Platemys radiolata is about the same size as platycephala but
is darker in color, the dorsal surfaces being uniformly black, and
has a deeper carapace with a less conspicuous median groove.
Also the eye, uniformly dark in platycephala, has a white iris in
radiolata. The head does not have the area of unbroken skin on the
crown so conspicuous in platycephala, the neck is covered with
pointed tubercles and the limbs with large flat scales. There is a
pair of barbels on the chin. The first and fifth central scutes are
very large, but the other three are much smaller and are about as
long as broad. Two subspecies (often considered as full species)
exist: radiolata from Brazil (Amazonas south to Sao Paulo) and
spixi from southern Brazil and northern Argentina. P. r. radiolata
has a light plastron with a large dark blotch nearly obscuring each
scute, while the plastron of spixi is uniformly dark.
780
Above: Plastral view of Platemys radiolata radiolata. Photo by Dr. Robert

Mertens. Below: The southern subspecies, Platemys radiolata spixi.
781
www.Ebook777.com
GENUS PHRYNOPS
SUBGENUS MESOCLEMMYS
Phrynops (Mesoclemmys) gibba, the only species of this sub
genus, is similar in coloration and size to Platemys radiolata but
has a median keel instead of a groove in the carapace. Neural
bones may be absent or there may be as many as five or six. This
condition is extremely unstable.
P. gibba has been recorded from Meta, Caqueta, and Vaupes,
Colombia; Para and Mato Grosso, Brazil; the Guianas; Trinidad;
Venezuela; eastern Ecuador; and northeastern Peru (Loreto). The
adult carapace length is about 6.5 inches; a 232.5mm specimen
from Ecuador is the largest recorded. The carapace is smooth and
keeled in all except the oldest specimens; the widest point of the
shell is somewhat behind the midpoint. The plastron is large and
has a prominent intergular separating the gulars and partially
separating the humerals. In color the plastron is brown with a
yellow border; the gulars are usually entirely yellow, and the
bridge yellow and dark brown. The submarginal area is yellow,
sometimes with a narrow brown border to each scute. The head is
of medium to large size, with a somewhat pointed snout. The iris of
the eye is uniformly pale yellow in color. The tympanum and lower
Widely distributed over northern and central South America, Phrynops

is a relatively small turtle with a narrow head.
(Mesoclemmys) gibba
jaw are cream-colored with small spots becoming denser and con
fluent on the throat. The top of the head is pale yellow marbled with
brown. Otherwise the skin of the soft parts is brown except for
yellow scales on the tip of the tail and ventral surfaces of the limbs.
The hind margin of the plastron is deeply indented, either roundly
or angularly. A specimen of this turtle from Caqueta, Colombia,
contained four eggs with hard shells; four other recorded clutches
contained 2, 3, 3, and 4 eggs respectively. Hatchlings are almost
two inches long. When picked up or otherwise disturbed, specimens
of this species emit a strong, distinctive odor.
This is a primarily nocturnal species, though it may bask in
the early morning or late afternoon. It is found both in still water
(muddy lagoons and ponds) and moving water (small forest
streams, small blackwater rivers, and streams running over a
muddy bed).
The remaining two groups of South American chelids, Phry

nops and Batrachemys, are closely related to each other and to
Mesoclemmys and are considered to be subgenera within the
genus Phrynops. Wermuth and Mertens distinguished Batra
chemys from Mesoclemmys by the shape of the rear emargination
of the plastron, but this character does not hold up with large
series. However, Mesoclemmys is a small animal, usually without
neural bones, and with only a moderately enlarged head; in adults
the width of the head is about one-sixth the length of the carapace.
The chin barbels are very small. Normally the intergular scute is
shorter than the sum of the interhumeral and interpectoral seams,
but a specimen from Parabam, Guyana (No. 61521 in the Ameri
can Museum of Natural History) has an intergular longer than the
sum of the two median seams behind it. Batrachemys is composed
of somewhat larger species a foot in length or more. Moreover
there are up to four neural bones (though sometimes there are
none) and the head is strikingly enlarged—about 1.75 inches wide in
the case of a seven-inch individual, while a seven-inch Mesoclem
mys has a head about 1 1/3 inches in width. The paired chin barbels
in Batrachemys are somewhat shorter than the diameter of the
eye. The temporal arch, connecting the parietal and squamosal
bones, is very slender, and the parietal bones present an extreme
ly narrow dorsal surface, so that the paired bunches of temporal
muscles are only separated by a ridge of bone. The species of
Phrynops range in length from ten to about eighteen inches and
have up to six neural bones. The head is intermediate in width be
tween those of Mesoclemmys and Batrachemys. The paired chin
barbels are sometimes very long, sometimes short, and the
783
www.Ebook777.com
parieto-squamosal arch is relatively strong. The parietal bones
present a moderately broad dorsal expansion. The systematics of
both of these subgenera are presently very poorly understood, and
the outline given below is likely to be altered in the future.
GENUS PHRYNOPS
SUBGENUS PHRYNOPS
One of the more distinctive species of Phrynops is P. rufipes. It

appears to be one of the rarest of all turtles, apart from certain
forms that have been depleted by man almost to the point of
extinction. Known specimens are as follows: the holotype, now
lost, collected by Spix in the Rio Solimoes (Upper Amazon) in 1820;
a male collected by Paul Mueller 28 km northeast of Manaus in the
Rio Negro in 1965; a female that I purchased from a dealer in
Florida in 1970, probably imported from the vicinity of Leticia,
Colombia; a live juvenile and female collected in Cano Colorado, a
tributary of the Pira-Parana in Vaupes, Colombia by the
anthropologists Stephen and Christine Hugh-Jones and Brian
Moser in 1970; and three empty shells from the same vicinity. By
far the most complete account of the species is Medem’s 1973
report; in this he summarizes the known range, from a
combination of actual specimens and reports by local Indians, as
comprising the area between the rivers Pira-Pirana, Apaporis,
and Papuri in Colombia, and the Upper Amazon, between
Tabatinga, Tefe, and Manaus in Brazil.
The most distinctive feature of Phrynops rufipes is the bright
red color of the skin of the head and limbs. This reflects in all of the
local Indian names—the Tucano name “U-suan”, the Bara name
“U-huwu”, the Barasana name “Gu-tsuwu”, and the Macuna
name “Musa-guga” all translate to “red turtle”. The carapace is
oval, with relatively large marginal scutes and a narrow, elongate
nuchal; in anterior profile the shell is strongly tectiform, and in
color it ranges from light brown in juveniles to dark brown in
adults. The plastron is yellow, slightly wider anteriorly than
posteriorly, and the abdominal scutes are distinctly foreshortened,
the interabdominal seam being the shortest along the entire
midline of the plastron. The hind lobe of the plastron has a deep
angular notch, and the intergular scute is typically large, widely
separating the gulars. However, two of the available specimens
show extensive abnormalities in the gular region, the intergular
being divided into three separate irregular scutes. Males have a
rather deep, angular plastral concavity, the form of which is
clearly adapted to accommodating the tectiform carapace of the
female during copulation.
The head of rufipes is broad and flattened, with a distinctly
protruding and pointed snout. The eyes are somewhat dorsally
784
situated, and the temporal region and cheeks are covered with a
mosaic of irregular scales; the skin on top of the head is smooth
and unbroken. There is a pair of very small barbels on the chin.
The red color of the head may be interrupted by a dark band
extending from the nostrils through the eye to the neck. The limbs
are long, with well webbed feet, and are well provided with ridges
and small scales. The claws are long and pointed and number five
on each front foot, but typically only three on each hind foot. The
maximum length is probably in the vicinity of nine inches.
According to the local Indians, rufipes is found in running
water in the rainforest; both ‘black’ and ‘white’ waters are
inhabited by this species. In diet it is primarily carnivorous; the
species may be caught on hooks baited with earthworms or insect
larvae. The stomach contents of the largest female reported by
Medem included remains of small fish and shrimp, as well as fruit
of the Assai palm (Euterpe). In captivity, Medem fed rufipes upon
raw meat. It is reported to nest between the beginning of July and
the end of August and to lay between 3 and 12 small, spherical,
hard-shelled eggs. The young first appear in September.
The Barasana and Tucano people do not consider rufipes to be
particularly rare, and indeed report it to be very common in the
Cano Uirari, below Araricuara. However, it is extremely shy and
not given to extensive basking so its presence would be easy to
overlook. This, combined with the restricted and very remote
range, would account for the very few specimens that have
reached the attention of herpetologists. In captivity it does not do
well; Medem’s specimens contracted a fungus disease and died
after a few months, which he attributed to the species’ need for
running water. My specimen was clearly diseased when I obtained
it, and it too was very short-lived in captivity.
Phrynops hogei was described by Robert Mertens in 1967 on
the basis of an adult male specimen, about 13.5 inches in length,
from the Rio Pequena near Sao Paulo, Brazil. The shell of this
holotype is smooth, low, unkeeled, and oval when viewed from
above. The first vertebral scute is very large, the second to fourth
rather small and slightly concave. The plastron is wider anteriorly
than posteriorly, is angularly emarginate behind, and shows a
very long interabdominal seam—twice as long as the interhumeral
or interanal seam. In P. rufipes, on the other hand, the interab
dominal seam is the shortest in the entire plastron. In color the
carapace—and indeed the entire dorsal surface of the animal—is
dark brown. The plastron is yellow-buff with dark infusions along
the sides, the bridges, and the underside of the marginals. The
head is very small for a Phrynops, about 1.8 inches in width, and
has smooth skin on the parts underlain by bone, but distinct granu-
785
www.Ebook777.com
lar scales on the temporal region. In dorsal outline the head is
more or less triangular in shape with a rather pointed snout. The
chin barbels are very small.
Phrynops hilarii is probably the largest member of the genus,
really big individuals approaching a carapace length of 18 inches.
This species is found in southern Brazil, Uruguay, and northern
Argentina. The carapace is parallel-sided, rather low, and with a
median flattening or slight depression in adults, a slight keel in
younger specimens. The front lobe of the plastron is broader than
the hind lobe; the suture between the pectoral scutes is much
shorter than that between the abdominals. The carapace is olive-
brown with a yellow margin; the plastron and submarginal area
are yellow with symmetrical black spots. The head is large and
very broad, with a narrow intense black line running from the
snout, through each eye, past the upper edge of the tympanum,
and along the neck. Above this line the top of the head and neck are
grey in color; below is almost immaculate white. The chin barbels
are very long and become broader from the root out. There is a
black ring around the base of each barbel, and in some specimens
a black line extends backward from behind each barbel to join the
Phrynops (Phrynops) geoffroanus from northern and central South America.

Specimen in the Frankfurt Zoo.
Head pattern of
Phrynops
(Phrynops) hilarii.
main head stripe at the back of the head. The upper surface of the
head is divided into irregular scales; the width of the skull at the
narrowest point across the parietal bones is about equal to the
diameter of the orbit. The inner surfaces of the limbs are cream or
yellow with one or more black spots or streaks; there is a black
band across the thigh region.
Phrynops geoffroanus is a slightly smaller species found from
Colombia through Venezuela, Guyana, and Brazil as far south as
Paraguay. In young specimens the carapace has a median keel
and reaches its widest point behind the midpoint. The hatchling is
very beautifully marked; the carapace is mottled with black and
grey-green, with pink markings at the extreme edge of each mar
ginal scute. The undersurface is pinkish-red with a complex pat
tern of irregular black blotches. The head is rather broad and flat,
the upper surface being black with irregular thin white lines, and
the undersurface white with black lines and spots; the most promi
nent head marking is a broad white line extending back and down
from behind the eye. The limbs are black with white markings. A
pair of long chin barbels is present. The attractive shell markings
fade with age but are still indicated even in adults. Two subspecies
have been described: geoffroanus from central and southern
Brazil and Paraguay, and tuberosus from Guyana and eastern
Brazil. It is possible that the Colombian population is referable to a
different subspecies, but no adequate comparison of specimens
has yet been made. The northern race, tuberosus, is recognizable
by the presence of three keels on the carapace (only one in geoffro
anus ).
787
www.Ebook777.com
Above: Phrynops (Phrynops) geoffroanus tuberosus. Photo by Dr. F. Medem.
Below: Phrynops (Phrynops) geoffroanus geoffroanus. Specimen from the
Columbus Zoo, Ohio.
788
In Colombia, this species is known as charapa and is found in

the Inirida, Vaupes, Apaporis, Caqueta, Putumayo, and Amazonas
systems; however, it is found with greater frequency in back
waters and lakes than in the rivers themselves. They bask in the
sun on half-submerged tree limbs or on the banks, and sometimes
basking aggregations of fifteen or twenty individuals are seen.
This species is carnivorous, eating small fish and insects. The
nesting season extends from December to February; the eggs are
hard-shelled, spherical, between ten and twenty in number, and
between 1 inch and 1.4 inches in diameter. The nests are made only
a few yards from the edge of the water.
Several other species, described under the names Platemys
wagleri, Hydraspis lutzi, and Hydraspis boulengeri, are probably
referable to Phrynops and may well be synonyms of forms de
scribed above; however, proper comparisons between adequate
series have yet to be made.
GENUS PHRYNOPS
SUBGENUS BATRACHEMYS
The taxonomy of the subgenus Batrachemys is also in a con
fused state. For many years Batrachemys nasuta was the only
species considered valid, but in 1958 Zangerl and Medem described
a new species from Colombia, B. dahli, and revived the species B.
tuberculata, originally described by Luederwaldt in 1926 from ma
terial from the Rio San Francisco in eastern Brazil. Then in 1969
Robert Mertens described a new chelid from northern Peru which
he called Phrynops wermuthi. However in 1970 Mertens wrote that
P. wermuthi was at best only subspecifically separable from
Batrachemys nasuta. The nomenclature we shall use here, then,
reflects the latest opinions but still must be regarded as provision
al.
Phrynops (Batrachemys) nasuta, the common toad-headed
turtle, is found over much of the Amazon drainage as well as in the
north-flowing river systems of the Guianas. The typical race,
P. (B.) n. nasuta, is a very large-headed turtle found in Brazil and
the Guianas. The carapace is mid-brown in color, rather low, often
flat-topped, and usually reaching its widest point behind the mid
point. The maximum shell length is one foot or a little more. The
plastron is rather large, slightly wider in front of the bridge than
behind it, and with an angular notch at the rear. The intergular
scute is shorter than the sum of the interhumeral and interpectoral
seams. The undersurface of the animal is basically yellow, but all
the plastral scutes except the gulars have extensive brown areas.
The snout is pointed, but the rear part of the head is very broad and
flat, with huge bulges of muscle for raising the lower jaw. The skin
789
www.Ebook777.com
tending the hind feet, so that the head and neck are less exposed to
attack. During the dry season they estivate, burying themselves in
soil or underbrush. In diet this species is primarily carnivorous,
eating snails, aquatic insects, tadpoles, frogs, small fishes, car
rion, etc., though some vegetable matter may also be eaten. There
is considerable sexual dimorphism, females reaching a length of
up to 8.5 inches, males not more than 6.75 inches. Males also have
narrower shells than females, with longer, thicker-based tails, and
tend to retain the head markings more clearly—the dark line run
ning from the nose to the neck remains distinct, the lateral parts of
the head and neck retain the yellowish juvenile color, and the red
dish ventral color also remains in adult males. Males moreover
have more strongly elevated and keeled scales on the latero-ven-
tral edge of the hind limbs. Mating occurs principally in June and
July, usually taking place in shallow water during the dawn and
dusk activity periods. The eggs number between one and six and
are hard-shelled and slightly ovoid; recorded maximum and mini
mum sizes are 35.5 x 29.0 and 28.0 x 23.5 mm.
Phrynops (Batrachemys) tuberculata (sometimes called
tuberculatus, the gender of Phrynops being debatable) is known
from eastern Brazil, having been recorded from the States of
Ceara, Bahia, and Pernambuco; Zangerl and Medem give the
range as the ‘basin of the Rio San Francisco.’ The species was
In Phrynops (Batrachemys) tuberculata from eastern Brazil the head is

uniformly grayish.
792
originally described by Luederwaldt in 1926 but was ignored for

more than three decades. It is distinguished from nasuta by the fol
lowing characters: i) the carapace reaches its widest point pos
teriorly rather than near the middle; ii) the middle three vertebral
scutes are narrower than in nasuta; iii) the last vertebral is 3 to 4
times wider behind than in front (about 2 times in nasuta)-, iv) the
nuchal is relatively wide; v) the dorsal keel is discontinuous and is
bordered by two gutters, which in old individuals coalesce posteri
orly to form a single median groove; vi) the intergular is longer
than its distance from the abdominals; vii) the inguinal is larger
than the axillary scute; viii) the width of the anal scutes is about
33% of the carapace length (narrower and with a deeper notch in
nasuta); ix) the limbs are relatively large and strong; x) the snout
is elongate rather than upturned; xi) the carapace is brownish-
yellow to blackish (red-brown in nasuta) -, xii) the head width is
about 24% of the carapace width (about 27% in nasuta); xiii) the
top of the head is greyish rather than brown or olive; xiv) the
lower jaw and throat are of the same color (contrasting colors in
nasuta). The habits of tuberculata have not been studied.
An old specimen of Phrynops (Batrachemys) vanderhaegei from Paraguay.

Photo by Dr. R.A. Mittermeier.
A new race of tuberculata, vanderhaegei, has recently been

described on the basis of an adult male specimen from Paraguay.
This for the time being might better be considered a full species,
since intergradation has not been demonstrated, and indeed the
ranges of the two forms, as presently understood, are widely sep
793
www.Ebook777.com
arated. Vanderhaegei differs from tuberculata in having no tuber
cles on the neck and in having a brown plastron. The holotype
lacks axillary scutes and has granular scales on the temporal
region. The supracaudal scutes are elongate and partially separa
ted by the posterior vertebral scute. The middle vertebral scutes
are narrow and show an interrupted dorsal keel. The vent is sur
rounded by a series of enlarged scales. The limbs are strong, as in
tuberculata; those parts which reach beyond the edge of the shell
are grey, while the more proximal parts are whitish. The jaws,
tympana, throat, and underside of the neck are uniformly pale yel
low. The otherwise grey dorsal surface of the head is marked with
brown or orange vermiculations. The wrinkles are somewhat
lighter than the surface. The brown iris is traversed by a horizon
tal black bar, though there are no indications of a continuation of
this dark bar on the skin.
AUSTRALASIAN CHELIDS
The Australasian genera of the Chelidae are four in number:
Chelodina, Emydura, Elseya, and Pseudemydura. Neural bones
are absent except in Chelodina oblonga. The genera may be con
veniently distinguished by the configuration of the intergular
scute: in Chelodina this scute is very large, completely separating
the humeral scutes and partially separating the gular and pectoral
scutes. The intergular does not border on the front edge of the plas
tron (except in an anomolous specimen of Chelodina siebenrocki
which was named as the type specimen of a new species, C. inter-
gularis). In Pseudemydura the intergular is also very large and
widely separates both gulars and humerals and partially sepa
rates the pectorals. Elseya and Emydura both have an intergular
which completely separates the gulars and partially separates the
humerals. These two genera may be distinguished by the single
horny shield covering the crown of the head in Elseya, which
develops at an early age and is present in all adults. The Austra
lian turtles, or tortoises as they are locally called, were, despite
their abundance and familiarity within the Australian continent,
very poorly known until recently. Several species were erected on
the basis of single anomalous specimens, and even the valid spe
cies were not well represented in museums, nor were they ade
quately described on the basis of large series of specimens. Now
an Australian herpetologist, John Goode, has published a book
entitled Freshwater Tortoises of Australia and New Guinea, which
represents a considerable advance in our knowledge of these tur
tles, though the New Guinea species remain very poorly known.
794
GENUS CHELODINA
Turtles of the genus Chelodina show an interesting parallelism
with the South American Hydromedusa—they have extraordinar
ily long necks, and in some species the length of the head and neck
approaches that of the carapace. The two genera, however, are
easily distinguished by the absence of neural bones (except in
oblonga) and the peculiar intergular scute of Chelodina, and the
wide, posteriorly-displaced nuchal scute of Hydromedusa. The
Australian snake-necked turtles, as the species of Chelodina are
usually called, have a very high frequency of scute abnormalities
in the carapace, specimens with six or more central scutes being
almost as common as those with five. The eight species currently
recognized are divided into three groups—the broad-plastroned
group (Chelodina longicollis, C. novaeguineae, and C. steindach-
neri) in which the length of the plastron is generally about 1.5
times the maximum plastral width (excluding bridge) and is
rarely if ever more than 1.75 X the plastral width; the narrow-
plastroned group (C. expansa, C. rugosa, C. parkeri, C. sieben
rocki), in which the plastron is at least 1.9X, and usually twice, as
long as broad; and C. oblonga, peculiar in possessing neural bones;
it has a narrow plastron.
Chelodina longicollis is by far the most familiar species. It is
also the most widely distributed, being found from Adelaide,
Mount Gambier, and Gippsland, in the south, throughout the Mur
ray River System, and along the eastern coastline as far north as
Rockhampton in Queensland, and inland in Queensland as far as
Stanthorpe. In some areas a shell length of about one foot is
reached, but adult specimens are usually about seven inches long.
The head is rather small, though it becomes somewhat larger in
old individuals, and the iris of the eye is brilliantly white and
staring. The shell is brown to black, with a shallow median groove
in mature specimens, and raised supracaudal scutes. In adult ex
amples the sides of the carapace curl upward in a distinctive
fashion. In texture the carapace is somewhat rugose; the outer
part of each scute is shed with growth, so ‘growth rings’ are not
normally present. However, occasional individuals do not desqua
mate in this fashion, and the distinctive appearance of such indi
viduals has led to their being described as a different species (sul-
cifera). But with the aid of a sharp fingernail, one can quickly con
vert a 'sulcifera' into a normal longicollis. The plastron is rather
broad, deeply notched behind and with blotchy lines of pigment
along the seams. The neck is dark dorsally and is covered with
small pointed tubercles. Murray River specimens have the widest
point of the carapace well behind the mid-point, and the hind edges
of the carapace are somewhat straight and meet at an obtuse
angle. In other areas (e.g. Gippsland) the shell is much more oval.
795
www.Ebook777.com
Juvenile Chelodina longicollis with the plastron mostly dark with pale blotch
es.
When picked up, fresh-caught specimens exude a strong-smelling

liquid from inguinal glands—a habit that has earned them the title
‘stinker’ in some areas. This species is known to feed upon aquatic
insects, frogs, tadpoles, and occasionally fish. Observations of
nesting habits are comparatively few, but in the Gippsland area
nesting is said to take place in November, 10-15 oval, hard-shelled
eggs being deposited at a time. Incubation periods of 130-137 days
have been recorded. Large numbers of animals may be prompted
to nest by heavy rain, and the nesting female also urinates in the
nest to assist with the digging process. Hatchlings are almost en
tirely black but have orange spots on each plastral shield and a
vivid orange stripe along each side of the lower jaw and neck.
Chelodina novaeguineae appears to be closely related to C.
longicollis, but differs in having blunt, rounded tubercles on the
neck, a somewhat narrower front plastral lobe, a totally unpig-
mented plastron, and a broader head. Also, the sides of the cara
pace are not curled up and the intergular scute is larger so that the
intergular and interpectoral seams are very short. The carapace
is typically somewhat lighter in color than that of longicollis, and
young specimens show very characteristic radiating striae on
796
each of the scutes of the carapace. The largest specimens are

about a foot in length, but the usual adult length is several inches
less than this. This species is widespread in New Guinea and is also
found in northern Australia, having been recorded from northwest
of Townsville, from the Cape York Peninsula, from Normanton,
and throughout the coasts of the Gulf of Carpentaria, from Arnhem
Land, and as far south as Katharine in the Northern Territory. It
has also been collected on the island of Roti, southwest of Timor. In
captivity it requires a much higher temperature than the very
cold-resistant longicollis. It has been found in swamps and
streams; its diet appears to be somewhat similar to that of C. long
icollis, but snails and aquatic vegetation are also eaten. It also
exudes a strong, unpleasant odor when molested. The egg clutch
size is said to range from 9 to 12.
Chelodina steindachneri is a relatively small species found in
rivers draining from the bulge of western Western Australia,
where it is the only turtle present. It has been recorded from as far
north as the De. Grey River and as far south as the Irwin River. It
is also known from the Murchison and Gascoyne Rivers and
probably occurs in the Fortescue, Ashburton, and Lyons Rivers. It
is found a considerable distance inland and is locally common as
far east as Wiluna, where A.A. Burbidge trapped eighteen
individuals in a 40-foot by 20-foot pool. The Canning Stock Route
The dinner-plate turtle,

Chelodina steindachneri.
Notice the pattern of
anterior plastral scutes
typical of most Australian
side-necks.
www.Ebook777.com
may represent the eastern edge of the range. To the north the
range is confined by the Great Sandy Desert and to the east by the
Gibson Desert. To the south, where the climate is moister and
cooler in winter, this species is replaced by C. oblonga.
C. steindachneri is commonly known as the plate-shelled or as
the dinner-plate tortoise, the names referring to the almost
discoidal carapace; Burbidge found the average shell is only 1.15
times longer than wide. The carapace is rugose and frequently has
a median groove. The largest female measured by Burbidge was
191 mm in length, the largest male 161 mm. The sexes are not easy
to distinguish, even in adults, although the female has a slightly
smaller tail. The plastron is broad, though much narrower than
the exceedingly broad carapace. The neck has dorsal reticulations
and is shorter and thinner than is typical of the genus, much more
so than the southern representative, C. oblonga. The color is
variable, some individuals have a light tan or pale olive shell, but
typically the shell has a bright red-brown color that matches the
water in which these turtles are found.
Water is scarce throughout the range of the dinner-plate
tortoise; some parts of the range have less than eight inches of
rain per year, while no part has more than ten inches. Such rain as
does fall usually comes all at once during the summer cyclones,
and many of the creeks in which the species is found only have
running water for a few days a year, drying up altogether for
months at a time. Temperatures may reach 40°C. or even 45°C.
The species is consequently better adapted for surviving arid
conditions than any other Australian turtle. It desiccates very
slowly, despite the lack of any obvious comification of the skin,
and can lose 38% of its body weight before it dies. It stores water
toward the end of the wet season in particularly capacious lateral
bladders; individuals are sometimes found wandering in parched
country a great distance from water, but they still expel a
remarkable volume of limpid water from the bladder when picked
up. This species has a well-developed ability to convert ammonia
to urates, which is of importance in detoxifying the urine without
having to discharge it.
Few data are available on the life history of the dinner-plate
tortoise, but local residents report that they lay their eggs in
spring (September to October) and that the eggs hatch when rain
begins in February. Hatchling emergence probably varies from
year to year, since rainfall is far from predictable. The
dinner-plate tortoise is assumed to estivate either in the sandy
bottoms of the dry creeks or under reeds and litter on the banks,
but this is unconfirmed. It may possibly seek refuge in burrows of
the local goanna lizard (Varanus gouldi). It feeds upon carrion and
also plant material, but, unlike C. oblonga, does not get big enough
798
to predate upon young aquatic birds. Burbidge caught these turtles

successfully in traps baited with the meat of the galah cockatoo,
but herpetologists in other continents are unlikely to emulate this
exotic technique. Burbidge has also found them easily caught with
a spotlight and a handnet at night, this method being particularly
suited to the small steep-sided pools in the intermittent rocky
creeks where this species is often found.
Chelodina millymillyensis, from Milly-Milly Station on the
Murchison River, is a synonym of C. steindachneri.
Chelodina expansa is the largest Australian chelid; maturity
is reached at a shell length of about eleven inches, and the shell is
known to reach a length of at least 16.5 inches. The range is said to
include inland Queensland and inland New South Wales to Murray
River and its tributaries in Victoria. The carapace is broadly oval
—slightly wider toward the rear and rather deep in large speci
mens. The plastron is very narrow. The head is large and very flat
and wide, and the neck is extremely long and does not have tuber
cles. The intergular is less than twice the length of the interpector
al seam. The supracaudal scutes are not raised. The carapace is
dark brown in color, the plastron anything from white to very
dark. There are two rudimentary barbels on the chin, or four in
some specimens. There are seven or eight lamellae on the upper
surface of each foreleg (usually four in longicollis and novae-
guineae). Goode writes that, when walking, this species holds its
head very low—below the level of the rim of the carapace. Chelo
dina expansa eats frogs, various aquatic insects, and, on occasion,
small fish. In the Murray River region of northern Victoria, nest
ing takes place from mid-March to May. The embryos overwinter
in the eggs and may take up to almost a year to hatch.
Chelodina oblonga is found in swamps and streams throughout
a wide area of the southwestern corner of Western Australia. The
carapace is elongate and very narrow, and becomes quite deep
with age. Most specimens have neural bones—in a series of ten
specimens, one had five neurals, three had six, four had seven, and
one had eight. The maximum carapace length is in the vicinity of
11 inches. The plastron is narrow, and the front lobe reaches its
widest point at the bridge. In texture the shell is rather smooth and
even glossy, bluish grey, brown, or black in color, and up to about a
foot in length. The neck is exceedingly long—longer, relatively,
than that of any other turtle species—and is so thick and muscular
that it is impossible for the turtle to draw it into the shell. The
hatchling has a posterior expansion of the carapace, but this is lost
with age. Rudimentary barbels are present on the chin. When
walking, the long neck is held straight in front of the shell, resemb
ling the front half of a slack-bodied snake. There are six or seven
lamellae on the upper surface of each foreleg. This species does
799
www.Ebook777.com
The neck of Chelodina obtonga of southwestern Western Australia is so thick
and muscular it cannot be completely withdrawn. Photo above of a specimen
in the Senckenberg Museum, that below in the Frankfurt Zoo.
800
not exude any odiferous material when molested. It is a popular

Item of native diet. Reports differ on whether or not the species
hibernates, and no data are available on dietary preferences or
nesting habits.
The species usually known as Chelodina siebenrocki is now
considered to be a composite of three species—Chelodina rugosa
from northern Australia (type locality: Cape York); Chelodina
siebenrocki from the southern coast of New Guinea and certain
islands in the Torres Strait, including Daru and Saibai; and Chelo
dina parkeri, newly described by Rhodin and Mittermeier from
inland southern New Guinea, apparently restricted to large inland
grass swamps surrounding Lakes Murray and Balimo and the Fly
and Aramia Rivers. A characteristic of members of this species
group is the egg-shaped or pear-shaped shell, very narrow in the
shoulder region but strongly expanded posteriorly. The neighbor
ing species siebenrocki and rugosa, clearly derived from a com
mon ancestor and separated by the Torres Strait, form a species
pair of a type equally exemplified by Elseya novaeguineae-E. den-
tata and Emydura subglobosa-E. kreffti. The populations of Chelo
dina novaeguineae north and south of the Torres Strait have
apparently not yet diverged morphologically.
Chelodina siebenrocki reaches a carapace length of at least
273 mm, and possibly as much as 380 mm. The carapace, although
strongly expanded behind, lacks flaring margins; the shell is
weakly domed, keelless, and the thin scutes have a rugose surface.
The rim between the upper skin of the neck and the shell margin is
very narrow, so the nuchal scute is considerably wider than long in
ventral aspect. The shell does not develop the central furrowing or
posterior median ridge shown by mature C. longicollis. The neck is
long and thick, and the head is flat, broad, and elongate, with a fine
reticulate pattern of skin creases which give the impression of
numerous small irregularly shaped scales. The chin barbels are
variable in number, but there may be four or even more. The plas
tron is much shorter than the carapace, and the anterior lobe is
very narrow. The intergular scute is almost twice as long as broad,
and in about 15% of specimens the intergular completely separates
the gulars and reaches right to the plastral margin (as in the type
specimen of the invalid Chelodina intergularis). The animal lacks
bright coloration, the dorsal surfaces being greyish-brown to black
and the ventral surfaces dirty white to light yellowish-brown. The
plastral seams may be finely outlined in dark brown. Males have
relatively flatter shells than females, and have longer, thicker
tails. Hatchlings have dark brown spots on the carapace.
Chelodina siebenrocki is apparently restricted to the tidal por
tions of the small streams flowing to New Guinea’s south coast and
to tidally influenced coastal swamps and offshore islands. It does
801
www.Ebook777.com
not bask, and spends most of its time in deep, soft mud. It is appar
ently carnivorous but not aggressive when handled; it is not
favored for food. Breeding appears to be seasonal, eggs being laid
at the end of the wet season in May, while hatching starts at the
beginning of the following wet season in November or December.
Two egg clutches counted by Fred Parker included four and seven
teen eggs respectively, the eggs being about 35-36 mm x 28-29 mm
in size. Hatchlings are 36-37 mm in carapace length. The species
occurs sympatrically with Chelodina novaeguineae, Emydura
subglobosa, Elseya novae guineae, and possibly also Carettochelys
insculpta and Pelochelys bibroni.
Detail of the head of Chelodina parkeri, formerly confused with Chelodina

siebenrocki. Photo by Dr. W. Sachsse.
Chelodina rugosa, the Australian relative of C. siebenrocki, is

still too poorly known for a complete description to be given. How
ever, the type specimen has a very narrow carapace and plastron,
while a series of seven Cape York specimens examined by Rhodin
and Mittermeier was too variable to permit conclusions to be
drawn. Much more work needs to be done on these turtles.
Chelodina parkeri is distinguished from C. siebenrocki by the
vermiculated color pattern of the head, the entire dorsal surface of
the head and the lower jaw being extensively vermiculated with
white, yellow, and pale green lines and dots on a dark brown back
ground. The pattern fades somewhat in adults, but always
remains. The shell is somewhat more parallel-sided than that of
802
siebenrocki; juveniles and subadults have vertebral tubercula-

tions. The intergular scute is not as elongate as that of siebenrocki,
and the carapace Is a lighter brown with pale yellow, indistinct
reticulations often present. The head is narrower than in sieben
rocki, and the chin barbels are smaller and fewer (usually only
two). The dorsal surface of the neck is covered with very small,
pointed, well-defined tubercles set closely together. The largest
specimen in the type series was a 267 mm female.
Chelodina parkeri is found in lakes and rivers in areas where
dense vegetation floats on shallow water; some of the situations
inhabited by this species are subject to partial or complete drying
during the dry season. The natural diet is unknown, but captive
specimens raised by Walter Sachsse fed exclusively upon fish. C.
parkeri shares its habitat with the much more abundant Emydura
subglobosa.
GENUS EMYDURA
The taxonomy of the Australian short-necked turtles of the
genus Emydura is also incompletely understood. Wermuth and
Mertens (1961) listed nine valid species: australis, albertisii, bran-
derhorsti, kreffti, macquarrii, subglobosa, latistemum, schultzei,
and novaeguineae. Worrell (1963) added a tenth species, E. vie-
toriae. However, Goode, in his 1967 book Freshwater Tortoises of
Australia and New Guinea, reduces the number of definitely valid
species of Emydura to three: macquarrii, kreffti, and australis. E.
latistemum and E. novaeguineae are placed in the genus Elseya;
branderhorsti, being based on a single specimen, poorly distinct,
and now lost, is relegated to the synonymy of E. novaeguineae, as
is E. schultzei which too is based on very inadequate material. The
New Guinea forms E. subglobosa and E. albertisii are provisional
ly synonymized with kreffti, pending further studies, and E. vie-
toriae is ignored. Goode’s designation of the turtles of the southern
and eastern parts of Australia can reasonably be accepted as
definitive, but it seems hasty to synonymize the northern and New
Guinean forms without concrete grounds for doing so.
The Murray River turtle, Emydura macquarrii, is the most
familiar species of the genus. Its range includes the length of the
Murray River and its tributaries from Adelaide, throughout
northern Victoria, and into inland New South Wales. Lower Creta
ceous fossil material found in Victoria was originally assigned to
this species in 1919, but the specimen has now been re-examined,
and has been declared the holotype of a new species, Chelycara-
pookus arcuatus, the only known species of the new family Chely-
carapookidae. Chelycarapookus is distinguished from E. mac
quarrii by having well developed posterior neural bones. Burbidge
suggests that it may be an early Australian chelid, arguing that the
803
www.Ebook777.com
The Murray River turtle, Emydura macquarrii macquarrii, of Victoria and New
South Wales.
presence of neurals in the living Chelodina oblonga suggests that

the original members of the family in Australia had those bones.
Genuine Emydura macquarrii fossils have been found in Oligo-
cene-Miocene formations in Tasmania. The shell is rather deep in
adults, with the highest point about one-third of the way back, and
is rather broad; in most areas the widest point of the shell is well
behind the midline, but in some areas (e.g. South Australia) the
carapace is almost circular. However, the shape of the carapace
alters with age; hatchlings are almost circular, with serrations on
the hind part of the carapace, half-grown specimens tend to be
widest behind, and large ones (the maximum length is about thir
teen inches) are almost oval. In texture the shell of large speci
mens is strongly rugose; in color the carapace is mid-brown, the
plastron yellowish. The head is rather small, with bright staring
eyes with yellow irises. A light band extends backward from the
comer of the mouth. There is usually a pair of barbels on the chin.
This is a highly active species, and when picked up it often strug
gles so hard that it is difficult to hold. The food is said to consist of
frogs, tadpoles, and aquatic vegetation. Up to 15 eggs are laid at a
time; they are of elongate shape, about 1*4 inches long x 7/8 inch
wide. Nesting takes place in northern Victoria from November
until mid-December, and the eggs hatch after about 10 or 11 weeks
under normal conditions.
804
A population of turtles of the genus Emydura found east of the

Great Dividing Range in Eastern New South Wales has been re
ferred to by the nomen nudum Emydura canni, but Dr. Robert
Mertens informs me that this population should properly be re
ferred to as Emydura macquarrii signata. My correspondent, Mr.
John Cann of Sydney, has sent me some interesting information on
these turtles. They differ from the forma typica of E. macquarrii
mainly in the smaller adult size (few of the over 2,500 specimens
examined by Mr. Cann exceeded a carapace length of 6% inches,
though the largest was eight inches long). The carapace may also
be darker and smoother, and the eyes a deeper yellow, especially
in the young. The range of this race is reported to extend north as
far as the Ebor River (30°25’S), and as far south as the Shoalhaven
River (approx. 35°S). The form is very abundant in the clear
waters of the Macleay River, where Mr. Cann has marked large
numbers of individuals and reported the interesting observation
that pairs of these turtles would swim together and remain in close
proximity to each other for periods of at least two months. The tur
tles showed a high degree of activity even when the water temper
ature was as low as 5°C., and were not in hibernation even at l°C.
Emydura kreffti is a northern relative of E. macquarrii; its
range extends from Cairns in the north to Kempsey (New South
Emydura macquarrii signata is a smaller, smoother version of the typical

subspecies.
www.Ebook777.com
Wales) in the south, being found only in east-flowing rivers and
their associated lagoons and swamps. Although somewhat similar
to E. macquarrii, kreffti may be recognized by the light bar exten
ding backward from the eye and the translucent carapace scutes
through which the bony sutures are clearly visible. The carapace
of the female is highly arched and reaches its highest point near
the mid-point. Males have a relatively flat shell. In both sexes the
shell is rather narrow. Barbels are not, or not usually, present on
the chin. A carapace length of about nine inches is reached. The
diet is said to include water snails, wild figs, and frogs. A specimen
in the possession of Mr. John Cann in Sydney laid 15 eggs on
December 2nd.
Emydura australis, locally known as the ‘boof-headed tor
toise,’ occurs throughout northern Australia from the Kimberleys
in the west to Katherine in the Northern Territory. The most ob
vious distinguishing feature of the species is the very large head of
the adult, with powerful jaws adapted for crushing molluscs,
though it also has been reported to eat berries, insects, and frogs.
The carapace is oval or only slightly expanded at the rear, and in
juveniles it has a serrated rear margin. The median keel of the
juvenile is lost with age. Young specimens lack the enormous head
of the adults and often have bright pink bands extending backward
from each eye and from each comer of the mouth, the latter
stripes often being continued along the neck. The pink color may
also extend to the shell and limbs, and the animal is so distinctive
at this stage that a separate name (Emydura victoriae) has been
applied to it. Emydura australis reaches a maximum carapace
length of at least l0.25 inches. According to a single observation
made by Eric Worrell in 1945, this species assumes a belly-to-belly
position when copulating. Neither the nesting procedure nor the
eggs have been described.
The precise status of the New Guinea species Emydura sub-
globosa and E. albertisii remains to be ascertained. It is probably
best to consider them as subspecies of australis. The late Prof. Dr.
Robert Mertens had in his private collection a handsome Emydura
from New Guinea which he considered to be albertisii, and which
certainly had a different color pattern from any Australian
species. This specimen was nearly a foot long and had a carapace
which was markedly expanded toward the rear. The plastral mar
gin and underside of the marginal scutes were suffused with bright
red. The limbs were black dorsally, white ventrally, and the head
was basically black but with a broad yellow band passing from the
snout through and above each eye and reaching to the back of the
head. The upper jaw was white, but the lower jaw was marked
with a prominent red streak on each side. There was no trace of
barbels on the chin.
806
Emydura australis
subglobosa, the
red-bellied side-
neck of southern
New Guinea and
the Jardine River
of Queensland.
GENUS ELSE YA
Turtles of the genus Elseya are distinguished from Emydura
by the following combination of characters: the narrow intergular
scute; a single horny scute covering the crown of the head; the
skin over the temples forming conspicuous flattened tubercles; tip
of the snout prominent; upper surface of the neck covered with
erect, pointed tubercles; prominent barbels always present on the
chin. Following Goode’s classification, three species are recog
nized: E. dentata, E. latisternum, and E. novaeguineae. On the
other hand, novaeguineae has recently been considered merely a
subspecies of latisternum. All are active, aggressive animals
whose habit of biting when handled has earned them the name
‘Australian snapping turtles.’
Elseya dentata occurs in rivers in inland northern Australia;
it has been found east as far as Hartley’s Creek near Cairns. The
type locality is the Victoria River. The carapace is slightly wider
posteriorly than anteriorly in all specimens; juveniles, however,
have a serrated rear carapace margin, while the entire shell rim
of the adult is smooth. The young have a median keel on the cara
pace, but in adults the shell is deep, rather flat-topped, and keel
less. In color the carapace is dark bluish-black, while the plastron,
unpigmented in juveniles, may become wholly black in later life.
The head is dark brown to black in young specimens, but with age
an increasing white area appears on the chin, throat, sides of the
head and neck, and also on the tip of the snout. This is the only Aus
tralian turtle with a median alveolar ridge. Adults reach a length
of at least thirteen inches; Worrell reports that the carapace may
anrr
www.Ebook777.com
Two views of Elseya dentata from northern Australia, perhaps the largest
chelid turtle. Photos of a specimen in the Senckenberg Museum.
808
exceed 18 inches in length, which would make this the largest

chelid turtle, though from the point of view of average adult size,
Chelodina expansa as well as one or two South American species
are larger. According to Worrell, Elseya dentata feeds on berries,
wild figs, water-weeds, water-lilies, crayfish, frogs, and some
small fish. It is a popular food item among aboriginals.
Detail of the head of

Elseya latisternum of
eastern Australia.
Elseya latisternum, commonly known as the saw-shelled

snapper, is found throughout the eastern coastline of Australia
from northern New South Wales in the south to Cape York; it is
also found in the Flinders River and possibly a considerable dis
tance around the Gulf of Carpentaria. Adults are easily recogniz
able by the strongly serrated hind margin of the carapace. The
shell is distinctively depressed and broad and may vary in color
from a rich chestnut in Cape York specimens to a somewhat
mottled dark and pale sepia from the Flinders River; some speci
mens are very dark brown. The maximum length is in excess of
one foot. The intergular scute is much wider than that of either
dentata or novaeguineae. The upper part of the neck is decorated
with a number of large, erect, pointed tubercles, and the chin bar
bels are also prominent. The snout is rather pointed. This is prob
ably the most aggressive Australian turtle, and it makes repeated
attempts to bite when molested. Worrell reports that this species
feeds upon frogs, crayfish, and aquatic vegetation. Despite its
wide range and local abundance, its nesting habits have not been
described.
The New Guinea snapper, Elseya novaeguineae, is widely dis
tributed on that island but does not appear to reach northern Aus
tralia. In this species the juvenile serrated carapace margin is lost
with maturity, but at least some indication of the vertebral keel is
809
www.Ebook777.com
Elseya novaeguineae is perhaps just a New Guinea subspecies of the
Australian Elseya latisternum.
retained. In young specimens the carapace is a rich chocolate-

brown, often with a single black blotch on each costal scute; the
carapace of the adult may be almost uniformly black. The shell is
deep and usually widest toward the rear; however, the holotype of
E. schultzei, which is commonly synonymized with this species,
has the widest point a little anterior to the mid-point of the cara
pace.
The eggs of E. novaeguineae, according to Nelly de Rooij, are
about 2.2 inches in length x 1.2 inches in width—much larger than
the eggs of, say, an Emydura macquarrii of comparable size. This
phenomenon—i.e. an increase in egg size (and presumably a de
crease in clutch size) from the temperate zone to the tropics—is
discussed in the survey of Rhinoclemys in the chapter on emydid
turtles.
GENUS PSEUDEMYDURA
The only remaining Australasian chelid is the diminutive and
rare Pseudemydura umbrina. This species is only found in the
Bullsbrook Swamp just north of Perth in Western Australia, and
has a curious history. The first specimen was collected in 1839 and
was deposited, without a name, in the Vienna Museum. It was not
810
until 1901 that Siebenrock described this specimen and proposed

the new genus Pseudemydura for it. No more were reported until
1954, when a specimen was received by the Perth Museum and was
described as a new species, Emydura inspectata. Finally, in 1958,
Ernest Williams compared this specimen with the type of Pseude
mydura umbrina and pronounced them identical.
This species has several points of interest. It is the smallest of
all chelid turtles, the record shell length being only 5.5 inches. The
carapace is wide and flat, and brown in color. The plastron is ex
tremely large, so that the tips of the anal scutes almost touch the
carapace. The intergular scute is heart-shaped and very big, com
pletely separating the gulars and the humerals and partially sepa
rating the pectorals. The plastral seams are dark-edged, and on
some specimens there are dark spots on several of the plastral
scutes. The head is very broad and flat and the snout flattened and
rounded. There are large homy tubercles on the neck and a pair of
small barbels on the chin. The skull is remarkable in that there is
very little emargination of the temporal region, either from below
or from behind. The limbs are unremarkable except for a unique
flap of skin on the hind margin of the forelegs, composed of three
or four enlarged scales.
The rare and endangered Pseudemydura umbrina, restricted to Bullsbrook

The Times.
Swamp, Western Australia. Photo by Dr. Ride and
811
www.Ebook777.com
Plastral view of Pseudemydura umbrina. Photo by Dr. Robert Mertens.
Because of the extremely restricted range and habitat prefer

ences of this turtle (it is only found in temporary swamps with a
clay or sand-over-clay bottom in the Upper Swan and Bullsbrook
area of southwestern Australia), it has been declared a threatened
species, and not only is it prohibited to collect specimens or import
the species into the United States, but the entire habitat (530 acres)
has been purchased for a reserve. The total population is calcu
lated at 200-300 individuals, and under this efficient protection,
over half of the turtles in the reserve at the present time are less
than three years old. It is known to feed upon tadpoles but does not
appear particularly adept at catching them. Three to five eggs are
laid at a time, in November or December; hatchlings emerge in
May or June. Six to nine years are required to reach maturity. A
captive group of about a dozen individuals is being maintained at
the South Perth Zoological Gardens, and some captive breeding
has taken place.
812
BACKGROUND READING
Ahl, E. 1931. Emydura signata sp. nov. Sber. Ges. Naturf. Freunde
Berl., 1931: 127-129.
Alarcon Pardo, H. 1969. Contribuction al comocimiento de la morfo-
logia, ecologla, comportamiento y distribucion geographica de
Podocnemis vogli, Testudinata (Pelomedusldae). Rev. Acad.
Colombiana de Clencias Exactas. Fis. y Nat. XIII, No. 51.
Barrett, C. 1929. Breeding notes on the Murray tortoise Emydura
macquarrii. Vic. Nat. 46: 95-96.
Bates, H.W. 1892. The naturalist on the Amazons. 5th. Ed., London.
2 vols.
Bour, R. 1973. Contribution a la connaissance de Phrynops nasutus
(Schweigger: 1812) et Phrynops tuberculatus (Luederwaldt:
1926). Description d’une nouvelle sous-espece originaire de
Paraguay, Phrynops tuberculatus vanderhaegei. (Testudinata-
Pleurodira-Chelidae). Bull. Soc. Zool. France 98(1): 175-190.
Budde, H. 1968. A rare Australian chelid? Int. Turt. Tort. Soc. J.
2(5): 12-14.
_________ , J.A.W. Kirsch, and A.R. Mair. 1974. Relationships
within the Chelidae (Testudines: Pleurodira) of Australia and
New Guinea. Copeia, 1974(2): 392-409.
Burbidge, A. A. 1967. The biology of South-Western Australian tor
toises. Ph.D. Thesis, University Western Australia, Perth.
Cole, M. and W. Link. 1972. Podocnemis in captivity. Int. Turt.
Tort. Soc. J. 6(2): 12-14, 33.
Curtiss, C. 1928. Notes on the egg laying of the long-necked tortoise
Chelodina longicollis. Qd. Nat. 6: 66-67.
de Rooij, N. 1915. Reptiles of the Indo-Australian Archipelago.
Brill, Leiden, pp. 314-320.
Dunn, E.R. 1945. Los generos de anfibios y reptiles de Colombia.
IV. Cuarta u ultima parte: Reptiles, ordenes Testudineos y
Crocodilineos. Caldasia, Bogota, 3(13): 307-335.
Ellenborgen, M. 1971. The Western Swamp Tortoise of Australia.
Int. Turt. Tort. Soc. J., 5(4): 16-17, 39.
Freiberg, M.A. 1971. El mundo de las tortugas. Editorial Albatros,
Buenos Aires. 134 pp.
__________ 1972. Validez especifica de Phrynops hilarii (D. et B.)
(Testudines, Chelidae). Act. Zool. Lilloana, XXIX: 247-260.
Fry, D.B. 1915. On a new Chelodina from Australia with a key to
the genus, Chelodina intergularis sp. nov. Proc. R. Soc.
Queensland, 27(4): 88-90.
813
www.Ebook777.com
Glauert, J. 1922. Chelodina milly-millyensis sp. nov. J. Proc. R.
Soc. West Aust. 9: 53-56.
Glauert, L. 1954. New swamp tortoise from the Swan River District
Emydura inspectata sp. nov. W. Aust. Nat. 4(6): 125.
Goeldi, E.A. 1889. Bedeutung, Fang und Verwertung der Schild-
kroten am Amazonas. Zool. Garten, Frankfurt am Main, 27:
329-436.
__________ 1906. Chelonios do Brasil (Jabotys, Cagados, Tartaru-
gas). Obra inedita, escripta entre 1892-1894. Bui. Mus. Goeldi,
Belem, 4: 699-756.
Goode, J. 1965. Freshwater tortoises in Victoria. Vic. Nat. 82:
169-175.
__________ 1967a. Freshwater tortoises of Australia and New
Guinea. Melbourne, pp. i-x, 1-154.
__________ 1967b. Artificial incubation of eggs of Victorian chelid
tortoises. Int. Turt. Tort. Soc. J. 1(3): 6-11, 41.
__________ 1968. Morphological variations. Int. Turt. Tort. Soc. J.
2( 2) : 8- 12.
Harrington, K.H. 1933. Breeding habits of Chelodina longicollis.
S. Aust. Nat. 15: 25-27.
Hartline, P. 1967. The unbelievable fringed turtle. Int. Turt. Tort.
Soc. J. 1(6): 24-29.
Hausmann, P. 1964. Nachzucht bei der Matamata. Aquar. und
Terrar. Zeit., 17(12): 369-372.
__________ 1968. Matamata. Int. Turt. Tort. Soc. J. 2(4): 18-19, 36.
Jacques, J. 1966. Some observations on the Cape terrapin. African
Wild Life 20(2): 137-150.
Kanberg, H. 1924. Uber eine neue Schildkrote aus Kamerun. Zool.
Anz., 60: 195-197.
Krefft, P. 1903. Die argentinische Schlangenhalsschildkrote. Bl.
Aquar. Terrar. Knde., Magdebourg, 14: 218-220, 228-230.
Laurent, R.F. 1956. Contribution a l’Herpetologie de la Region des
Grands Lacs de l’Afrique centrale. I. Generalites. II. Chelo-
niens. III. Ophidiens. Ann. Mus. Roy. Congo Beige, Zool., 48,
pp. 1-190.
__________ 1964. Batraciens et reptiles de l’Angola. Troisieme
Note. Publ. Cultur. Comp. Diamantes Angola, 67, pp. 1-165.
__________ 1965. A contribution to the knowledge of the genus Pelu-
sios (Wagler). Mus. Roy. de l’Afrique Centrale, Tervuren, Ser.
8 (Sci. Zool.), no. 135: 1-33.
Loveridge, A. 1941. Revision of the African terrapin of the family
Pelomedusidae. Bull. Mus. Comp. Zool., 88: 467-524.
Luederwaldt, H. 1926. Os chelonios brasileiros com a lista dos
especies do Museu Paulista. Rev. Mus. Paulista, Sao Paulo, 14:
405-470.
814
Medem, F. 1960a. Informe sobre reptiles colombianos (IV). El hal-

lazgo de la tortuga Phrynops (Batrachemys) nasuta (Schweig-
er) en Colombia. Nov. Colom., 1(5): 284-290. Popayan.
_____ . 1960b. Datos zoo-geographicos y ecologicos sobre los
Crocodylia y Testudinata de los Rios Amazonas, Putumayo, y
Caqueta. Caldasia, VIII 38: 341-351.
__________ 1964. Morphologie, Okologie and Verbreitung der
Schildkrote Podocnemis unifilis in Kolumbien (Testudinata,
Pelomedusidae). Senckenbergiana biol. 45(3/5): 353-368.
__________ 1966. Contribuciones al conocimiento sobre la ecologia
y distribucion geographica de Phrynops (Batrachemys) dahli;
(Testudinata, Pleurodira, Chelidae). Caldasia IX (45): 467-489.
______ . 1973. El primer hallazgo de la tortuga Phrynops rufipes
(Spix) en Colombia. Revista de la Academia Colombia de
Ciencas, 4:49-68.
Mehrtens, J. 1967. Chelids of Australasia. Int. Turt. Tort. Soc. J.
1(6): 14-21.
Mertens, R. 1967. Bemerkenswerte Susswasserschildkroten aus
Brasilien. Senckenbergiana biol. 48(1): 71-82.
__________. 1969a. Bemerkungen uber einige Halswender-Schild-
kroten. Senckenbergiana biol. 50(1/2): 31-38.
__________ 1969b. Eine neue Halswender-Schildkrote aus Peru.
Senckenbergiana biol. 50(3/4): 132.
__________ 1970. Zur Kenntnis von Phrynops nasutus. Senckenber
giana biol. 51(1/2): 17-20.
Mittermeier, R.A. 1975. A turtle in every pot. Animal Kingdom,
April/May:9-14.
__________, F. Medem, P. Soini, M.S. Hoogmoed, and A.G.J.
Rhodin. In Press. Distribution of the South American chelid
turtle Phymops gibbus, with notes on habitat and repro
duction. Herpetologica.
______ ___ and R.A. Wilson. 1975. Redescription of Podocnemis
erythrocephala (Spix, 1824), an Amazonian pelomedusid
turtle. Papeis Avulsos de Zoologia, Sao Paulo, 28(8): 147-162.
Mondolfi, E. 1955. Anotaciones sobre la biologia de tres Quelonios
de los Llanos de Venezuela. Mem. Soc. Cis. nat. La Salle, Cara
cas, 15(42): 177-183.
Muller, L. 1935. uber eine neue Podocnemis-Art (Podocnemis
vogli) aus Venezuela nebst erganzanden Bemerkungen uber
die systematischen Merkmale der ihr nachstverwandten
Arten. Zool. Anz., 110: 97-109.
_________ . 1939. Uber die Verbreitung der Chelonier auf dem suda-
merikanischen Kontinent. Physis, Buenos Aires, 16: 89-102.
Muller, P. 1966. Ein Wiederfund der Roten Krotenkopf-Schildkrote
(Phrynops rufipes). Aquar. Terrar. Z. Stuttgart, 19: 373-374.
815
www.Ebook777.com
Neill, W.T. 1965. Notes on the five Amazonian species of Podoc-
nemis (Testudinata, Pelomedusidae). Herpetologica 21: 287-
297.
Niceforo Maria, Hno. 1952. Testudineos del Suborder Pleurodira en
el Museo de La Salle. Bol. Inst. La Salle 39: 1-8. Bogota.
Oliver, J.A. 1959. The matamata—surely all turtles can swim. . .?
Animal Kingdom 62: 167-170.
Parsons, J.J. 1962. The green turtle and man. Univ. of Florida
Press. 125 pp.
Ramirez, M.V. 1956. Estudio biologico de la tortuga ’arrau del Ori
noco, Venezuela. Agricultor Venezolano 21(190): 44-63.
Roze, J.A. 1964. Pilgrim of the river. Nat. Hist., N.Y. 73(7): 34-41.
Schmidt, A.A. 1966. Morphologische Unterschiede bei Chelus fim-
briatus verschiedener Herkunft. Salamandra, 2(3): 74-78.
Schmidt, K.P. 1919. Contributions to the Herpetology of the
Belgian Congo based on the Collection of the American Museum
Congo Expedition 1909-1915. Part 1. Turtles, Crocodiles, Lizards
and Chameleons. Bull. Amer. Mus. Nat. Hist., 39: 385-624.
Schnee, P. 1907. Eine seltsame Schildkrotenart (Platemys spixii
Dumeril und Blbron). Wschr. Terrar. Kunde, Braunschweig,
4: 195-196.
Siebenrock, F. 1901. Pseudemydura umbrina sp. nov. Beschrei-
bung einer neuen Schildkrotengattung aus der Familie Chely-
didae von Australien: Pseudemydura. Sber. Anz. Akad. Wiss.
Wien 22 : 248.
__________ 1902. Zur Systematik der Schildkroten-Gattung Podoc-
nemis Wagl. S.-B. Akad. Wiss., math-naturw. Cl., Wien, 111:
157-170.
__________ 1904. Schildkroten von Brasilien. Denkschr. math.
naturw. Kl. Akad. Wiss. Wien, 76: 1-28.
__________ 1905. Uber die Berechtigung der Selbstandigkeit von
Hydraspis hilairi Dumeril und Bibron. Zool. Anz., Leipzig, 29:
424-427.
__________ 1906. Schildkroten von Ostafrika und Madagascar. In
Voeltzkow, 1906-1910, “Reise in Ostafrika in den Jahren 1903-
1905.” 2, pp. 1-40.
__________. 1909. Synopsis der rezenten Schildkroten mit Beruch-
sichtigung der in historischer Zeit ausgestorbenen. Zool. Jb.
Supp. 10: 581-586.
__________ 1916. Schildkroten aus dem nordlichen Seengebiet und
von Belgisch Kongo. Ann. Naturhist. Hofmus. Wien, 30: 1-12.
Silva Coutinho, J.M. da. 1868. Sur les tortues de l’Amazone.
Traduit du Portugais, sur manuscrit inedit. Bull. Soc. imper.
Zool. Acclim., Paris, (2)5: 147-166.
Smith, N. J.H. 1975. Destructive exploitation of the South American
river turtle. Chelonia, 2(5): 3-9.
816
Stejneger, L. 1909. Generic names of some chelid turtles. Proc.

Biol. Soc. Washington, 22: 125-128.
Venezuelan Embassy, Washington. 1958. Arrival of turtles in har
vest time for fishermen. Venezuela up-to-date, Feb. 1958, p. 17.
Vogt, T. 1911. Emydura schultzei sp. nov. Reptilien und Amphibien
aus Neu Guinea. Sber. ges. naturf. Freunde, Berl. 9: 410-412.
Waters, J. 1972. Survivors of ages past. Int. Turt. Tort. Soc. J.
6(4): 28-34.
Wermuth, H. and R. Mertens. 1961. Schildkroten, Krokodile, Bruc-
kenechsen. Fischer, Jena. 422 pp.
Williams. E.E. 1954. A key and description of the living species of
the genus Podocnemis (sensu Boulenger), (Testudines, Pelo-
medusidae). Bull. Mus. Comp. Zool., 111(8): 279-295.
__________ 1954b. A new Miocene species of Pelusios and the
evolution of that genus. Breviora, No. 25: 7 pp.
__________ 1958. Rediscovery of the Australian chelid genus Pseu
demydura Siebenrock. Breviora (84): 1-9.
Wood, R.C. 1973. A possible correlation between the ecology of liv
ing African pelomedusid turtles and their relative abundance
in the fossil record. Copeia, 1973(3): 627-629.
__________ 1974. The systematics, ecology, evolution, and zoogeo
graphy of African turtles. Natl. Geog. Soc. Res. Reports, 1967
Projects: pp. 301-306.
Worrell, E. 1945-46. E. australis “Boof head” tortoise eating fresh
water mussels (Propepydridellidae). Proc. Zool. Soc. London,
pp. 29-31.
__________ 1961. C. oblonga synonymous with C. intergularis. W.
Aust. Nat. 8: 18-27.
__________ 1963. Reptiles of Australia. Angus and Robertson, Syd
ney. Pp. 11-17.
Zangerl, R. 1948. The vertebrate fauna of the Selma Formation of
Alabama. Part II. The Pleurodiran Turtles. Fieldiana Geol.
Mem., 3: 3-56.
__________and F. Medem. 1958. A new species of chelid turtle,
Phrynops (Batrachemys) dahli, from Colombia. Bull. Mus.
Comp. Zool., 119 : 375-390.
817
www.Ebook777.com
Chapter 11
TURTLE CONSERVATION
AND EXPLOITATION
Only a few years ago, concern for vanishing species and dim
inishing natural resources was the monopoly of a select group of
mostly privileged people. Now that the results of years of environ
mental abuse are apparent on all sides, the topic of ‘conservation’
is familiar, in some degree, to nearly everyone. Conservation liter
ally means the act of conserving, or maintaining, something in its
original state, abundance, or extent, and it is quite legitimate to
speak of conserving water, green turtles, wilderness, the Okla-
waha River, forest land, and many other diverse natural products
and phenomena. The strict definition of conservation does not in
clude complete protection, as opposed to rational use, of a particu
lar species, though the practical conservationist may often find
himself recommending that a rare species should be completely
protected, either because it has been over-exploited in the past or
because its natural range is so restricted and its population dy
namics so poorly known that any exploitation may be hazardous to
the survival of the species.
The desirability of conservation of a natural resource is ob
vious to anyone who is able to think beyond his immediate needs.
However, when one’s immediate needs are strong it is often diffi
cult to see beyond them, and in many cases when a single resource
is being exploited by a variety of individuals—or nations—the atti
tude is adopted that ‘if we don’t get it now, someone else will get it,
either now or later,’ and the result is a mad scramble to get what
one can while one can. It also happens that people are on the edge
of starvation, leading a hand-to-mouth existence, in which case
laws protecting edible animals are a nicety which they cannot
afford to follow. Others—the most selfish of all—simply say that
they will take all they can now, and when the resource becomes ex
tinct, they will turn to something else. Many exploiters—ranging
from individual turtle poachers in the Caribbean to the Japanese
whaling industry—exemplify this attitude.
Despite these exceptions, the desirability of conservation, at
least in abstract terms, is evident to most people if it doesn't entail
any sacrifice on their part. The desirability of protection, however,
is less widely recognized, since its benefits are less concrete. In
818
many cases protection involves a definite inconvenience to people,

or restriction of their activities, and may cost a lot of money and
effort without any certainty or even strong likelihood of direct
material return for one’s investment. Sometimes we can talk in
terms of indirect material return; a rare species, well protected in
an attractive natural habitat, may attract paying tourists and
sightseers. However, one’s ultimate arguments for protection
even of obscure, unspectacular, rare, and materially useless ani
mals involve respect for the right of other organisms to have their
own place on the face of the earth, respect for one’s environment,
harmony with nature, and esthetic appreciation of the diversity
of the forms of life—arguments which, to many people, are mean
ingless. Some argue that it is part of the natural scheme of things
for species to become extinct, and that it is misplaced sentimen
tality to try and preserve them. The fault in this argument is that
the present rate of extinction—hundreds of vertebrate species
every century—is not natural, since the species are not being re
placed by newly-evolved forms, and the net result is a lowering of
the diversity of existing life. Man has the power to exterminate
many species which are perfectly adapted to their environment
and which are not in any way inept or obsolete. If by some miracle
a population of dinosaurs was to be discovered in some remote
part of the world, there would be strong—and justified—clamor to
keep them alive, despite the fact that this form of life has been
superceded elsewhere by organisms better adapted to present con
ditions.
Turtles rarely make a nuisance of themselves, except perhaps
to the ornamental horticulturist whose flower beds are dug up by
nesting snapping turtles or fishermen who get annoyed because
they keep catching mud turtles instead of fish, and while many
people remain unconvinced that crocodiles, venomous snakes, and
some other reptiles have a right to live, they are not on principle
averse to sharing the earth with a few turtles. Nevertheless it is
probably true that every species of turtle has become reduced in
numbers to some extent by man, and the population decline of
some species has been catastrophic. Turtles are destroyed, either
as individuals or as entire populations, by a variety of human acti
vities. The most obvious of these is direct consumption of turtle
meat and eggs; with very few exceptions all turtles are edible, and
frequently they are found in places where the human population is
undernourished and protein-starved. Turtle steaks and soup
(made from the green turtle) are also consumed by the affluent in
the United States and western Europe. Certain species of turtle are
killed for non-food products—the ridleys for their skins (for luxury
leather) and hawksbills for their shells, the plates of which are
removed and polished for ‘tortoiseshell.’ The pet trade removes
many turtles from the population, and in recent years a tremen-
819
www.Ebook777.com
dous variety of unusual and exotic species has been exploited for
this purpose, although in the last couple of years increasing re
striction on the export and import of exotic animals is reducing the
variety available to turtle fanciers, as well as increasing prices.
Turtles are killed for scientific study, for museum collections, for
class dissection, and for neurological studies. Many turtles are
killed by vandals, as well as by sports fishermen who mistakenly
believe that turtles feed on game fish or who get frustrated by the
frequency with which they catch turtles, rather than fish, on their
lines.
Whole populations of turtles can be wiped out in an indirect but
highly effective manner when their habitat is upset or destroyed;
drainage of lakes and swamps, introduction of foreign predators or
competitors, extermination of food organisms, and land clearance
schemes have all played their part in the reduction of turtle popu
lations.
It is worthwhile discussing in greater detail each of these
causes of loss of turtles and evaluating which are defensible, which
avoidable, and which unnecessary.
TURTLES AS FOOD
In many parts of the world—Europe, most of Australia, the
Arab countries, and parts of Southeast Asia—turtles are rarely
eaten, except in the form of green turtle soup, which is highly es
teemed as a luxury item by Europeans. The bases for this reluc
tance to eat turtles are not always readily discernible, though
where standards of living are high and turtles few, it is natural
that turtles should be regarded with esthetic rather than gusta
tory appreciation. Religious beliefs prohibit eating of turtles in
many parts of the Arab countries, Israel, and parts of Southeast
Asia, though often turtle eggs—particularly sea turtle eggs—re
main a legitimate and in fact over-exploited commodity.
True tortoises of the family Testudinidae have a unique and,
for them, disastrous combination of characteristics; they are the
only animals that are at the same time excellent to eat, ridiculous
ly easy to catch, and with virtually no way of defending themselves
when picked up, except by withdrawing into the shell and thereby
making themselves a very convenient bundle to carry. Moreover
they have a droll demeanor that commends them as pets to those
who are sufficiently well-nourished not to have to consider them as
food, though they do not do particularly well in captivity, especial
ly when kept indoors. Migrant shepherds in Morocco sell all tor
toises they find to collecting stations, where they are either packed
in baskets and shipped alive to Europe, or those that are too small
or too large for the legal pet trade are killed and their shells made
into banjos. The tortoise populations of Morocco have apparently
820
been reduced in many areas, though larger numbers remain in

Algeria and Tunisia.
Throughout much of tropical Africa it is probable that the lar
ger tortoises (in particular the leopard and spurred tortoises) are
eaten whenever they are found, while Kalahari bushmen will even
eat the diminutive serrated tortoise (Psammobates). In Madagas
car the species Geochelone yniphora has apparently been almost
exterminated, and Robert Baudy reports that the beautiful radia
ted tortoise, Geochelone radiata, is now disappearing fast; al
though it is fully protected by law and is not eaten by the natives in
the area in extreme southern Madagascar to which it is now re
stricted, Chinese in Tananarive will pay the equivalent of $50 for a
tortoise for eating purposes, and a lively illegal trade in the species
exists. Double-bottomed pick-up trucks are used to smuggle the
tortoises to the capital. In view of the probable extinction of the
species in the wild, Robert Baudy obtained permits to export sev
eral dozen radiated tortoises to his farm in Center Hill, Florida,
which offers a reasonable facsimile of the climate to which the tor
toises are exposed in the wild. All of the tortoises grew substantial
ly when set up in their new quarters, and as of the time of writing
Tortoises such as this red-footed tortoise, Geochelone carbonaria, have long

been used as both food and pets. Photo by R.J. Church.
www.Ebook777.com
only one death has occurred. This colony has now been transferred
to the care of Dr. Walter Auffenberg in Gainesville, and some
reproduction has taken place.
Throughout tropical lowland South America, the two native
tortoises (the red-foot and the yellow-foot) are eaten whenever
found, as is Rhinoclemys in northern South America. However,
individuals of these species are widely dispersed, and at existing
human population levels ample reserves remain in the huge and
sparsely inhabited forest areas, though around human settlements
tortoises disappear rapidly. Much more vulnerable are the aquatic
Podocnemis species (especially P. expansa) which nest colonially
on sandbanks in the Orinoco and Amazon River systems. For cen
turies these turtles have been exploited for meat, oil, and eggs on
their nesting grounds, and now the populations are seriously deple
ted. A few years ago pet shops in the United States sold bulk quan
tities of hatchling Podocnemis expansa and unifilis, but now
importation of these two species into the United States without a
federal permit is prohibited. P. expansa is now protected in much
of Brazil and is being actively conserved in Venezuela, where
hatchling tagging and re-introduction programs have been
started.
col
Peltocephalus tracaxa
lected for the market. Photo
by Federico Medem.
In Central America exploitation of freshwater turtles is desul

tory, but in many areas sea turtles and their products are items of
large-scale commerce. This industry is discussed later. In Mexico
freshwater turtles are eaten in quantities in the southeastern
states; Dermatemys, Pseudemys, Staurotypus, Chelydra, Kino
sternon, and Claudius are the most-used forms. Dermatemys is
theoretically protected during certain months. Further north, the
rare Bolson tortoise, Gopherus flavomarginatus, is prized for its
flesh, and its small range is diminishing each year. Although it has
822
been reported that the local people set out food for the tortoises in
the dry season and only eat a carefully limited number of the tor
toises each year, Bruce Deuley reports that some ranchers in the
area kill all tortoises they find because they compete with cattle
for grass.
In the United States, the diamondback terrapin was highly
favored several decades ago and farms were established to protect
the species from extinction. Now diamondbacks are relatively
rarely eaten and are again common in many areas, though it is
unlikely that the farms had anything to do with this comeback.
Snapping turtle was and to some extent still is a traditional favo
rite in Philadelphia, but the number consumed does not appear to
be excessive in view of the abundance and wid 3 range of the spe
cies. In the Southeast, soft-shells and Suwanr^e cooters are the
preferred forms. Some black people in the s ,'uth have a great
appetite for gopher tortoises, and for this reason, as well as wide
spread habitat destruction and numerous road kills, a substantial
reduction in gopher populations has occurred during the last ten
years. Gopher tortoises are not protected in Florida, though recent
legislation prohibits the gopher, together with the alligator snap
ping turtle and Barbour’s map turtle, from being bought or sold.
Sliders have been exploited tor food and as laboratory animals but are still
abundant. Photo of Pseudemys concinna concinna.
www.Ebook777.com
The Texas and desert tortoises are both protected by law, but large
numbers are still being diverted to the pet industry, and I am in
formed that Texas tortoises are also exported from the state ille
gally for the cosmetics industry.
The turtles whose existence has been most seriously threat
ened by the demand for their flesh are the big ones—the Indian
Ocean and Galapagos giant tortoises and the sea turtles. All but
one of the Indian Ocean forms have been extinct since before 1800
as a result of a combination of massive over-exploitation and intro
duction of domestic animals, especially pigs, to their habitat. Only
the Aldabra tortoises survive, but they are very abundant—esti
mates for the population on the 18-mile long atoll range from 30,000
to 100,000. The only conservation measures required at the present
time are control of exports of live animals to a fixed and very low
annual quota (the species is no longer on the AAZPA threatened
list, and the proprietor of the island presently exports 50 tortoises
annually), and to ensure that no habitat changes occur—in par
ticular, to ensure that the postponed establishment of a British air
base on the island never materializes.
All the species of sea turtles are presently listed in the Red
Data Boole—the official IUCN list of threatened species. This is not
because existing populations are already reduced to a low absolute
level; the green turtles of the world, for example, may still num
ber in the millions. However the rate of decline of some species is
distressing, and all are exceedingly vulnerable when the females
emerge from the sea to lay their eggs. Active conservation of sea
turtles, concentrating on the economically very important green
turtle, was initiated in the 1950’s by Archie Carr at Tortuguero,
Costa Rica, and by Tom Harrisson on the Talang Islands of Sara
wak. The methods of conservation employed in the two areas were
very different, reflecting the great differences in the local patterns
of exploitation. In Costa Rica the main problem was the excessive
numbers of turtles that were turned over on the beaches for later
collection and slaughter. Laws were passed and beach patrols con
ducted throughout the nesting season, whose effect has been to re
duce considerably, though not to eliminate, the killing of nesting
turtles. Recently the zone in which turtles could not be taken at sea
was extended from one to five kilometers from the beach, but an
agreement not to exploit green turtles drawn up by Costa Rica,
Nicaragua, and Panama (the latter two countries being the princi
pal feeding areas for the majority of greens nesting at Tortuguero)
fell through. Visible benefits from this beach protection are
masked by the astonishing year to year fluctuations in the num
bers of turtles nesting at Tortuguero. For example, 1968 was the
worst year on record, while 1969 was the best, and 1970 was an
average to good year.
824
Tortuguero has also provided a source of hatchling turtles for

‘Operation Green Turtle.’ This program, initiated by Archie Carr
and assisted by the United States Navy, involved the transporta
tion by air of tens of thousands of green turtle hatchlings from Tor
tuguero to a number of Caribbean islands and territories which
had formerly provided nesting grounds for the species. The results
were ambiguous; although turtles were transported in each season
from 1957 to 1968, no new nesting colonies are known to have been
established. On the other hand, healthy turtles of near-adult size
are now being found in areas where they had not been seen for
many decades.
Tom Harrisson’s work in Sarawak was made simpler by the
fact that the local people, for cultural reasons, never ate turtle
meat or killed turtles for any purpose. Perhaps with an intuitive
grasp of the rather non-dynamic population structure of sea tur
tles, they elected not to kill the goose that laid the golden eggs. The
golden eggs, on the other hand, were collected on a total exploita
tion basis, and the necessity was to arrange to have set aside a pro
portion of the eggs for hatching. In recent years the Sarawak green
turtle colony has shown a marked downward trend. This may be
because too few eggs were set aside for hatching, but may also be a
result of killing of the adult turtles in the Philippines or in the open
sea.
A few years ago the governments of Surinam, South America,
and Queensland, Australia, totally prohibited the killing of turtles
or digging up of their eggs. Enforcement is good, and the future for
turtles in these two territories will be as secure as it is anywhere as
long as present government policies are maintained. In each case
the prime moving force alerting the governments to their conser
vation responsibilities has been one man—Johan Schulz in Suri
nam and Robert Bustard in Australia. George Hughes of Durban,
South Africa, has also been successful in initiating effective regu
lations and patrols for protecting nesting leatherbacks and logger-
heads in Natal. The State of Sabah, in Malaysian Borneo, has an
effective program for conserving green turtles and hawksbills
under the direction of the Chief Game Warden, Stanley de Silva,
and the government of Trengganu, Malaysia, buys back a small
percentage of leatherback eggs laid on its beaches for artificial in
cubation. The proportion set aside is very small (2-5%), but it is
hoped that it will be increased in the near future.
In Mexico the government endeavors to protect turtle eggs
and hatchlings for largely commercial reasons, and it is compul
sory for shelled eggs taken from turtles, even when legally caught
at sea, to be buried, and the percentage hatch reported to the fed
eral government, which issues a leaflet giving instructions on arti
ficial incubation of turtle eggs. Unfortunately, the coastline of
Mexico is so long that enforcement of conservation laws is loca-
825
www.Ebook777.com
lized and sporadic, and the number of adult turtles killed, legally
and illegally, is probably greatly excessive. An 18-month closed
season on all turtle hunting in Mexico recently expired, and heavy
exploitation is again in progress.
In the British Indian Ocean Territories green turtles are now
seriously depleted and, though presently protected by Seychelles
law, moves are afoot to reverse the existing legal protection. Ex
ploitation of the huge nesting colony to the north, in South Yemen
and Pakistan, appears to be desultory, civil strife having put the
exporters out of business.
In the United States molestation of nesting turtles or their eggs
carries a heavy fine, and in many areas enforcement is quite good,
though a loophole in the law is that turtles may be killed as soon as
their flippers touch water on their return to the sea. The danger to
the American populations lies more in beach habitat destruction
and the activities of shrimp boats. In Hawaii, however, any turtle
found near the inhabited islands is killed for food, turtle meat
fetching a very high price in Honolulu restaurants.
A novel development of great relevance to green turtle conser
vation is the establishment of a highly sophisticated turtle farm,
operated by Mariculture Ltd. with Anglo-American financing* and
leadership, on Grand Cayman Island. The object of the enterprise
is to produce a dependable and competitively-priced supply of tur
tles to satisfy the luxury market. As yet the operation is still depen
dent of eggs brought from natural nests in Costa Rica, Ascension,
Guyana, and Surinam, but it is to be hoped that ultimately all tur
tles needed will be bred from captive stock. A significant step in
this direction is the successful breeding by adult turtles, flown in in
gravid condition from Costa Rica, on an artificial beach at one side
of a large tank excavated out of the rock. In 1973 approximately
12,000 eggs were laid in this beach. If all succeeds as planned,
Mariculture could well be of great benefit to turtle conservation by
supplying the luxury demand entirely from farm-raised animals.
On the other hand, Mariculture’s egg-collecting activities on a
number of beaches on which local people are forbidden to collect
eggs is a most unfortunate example, and also it is conceivable that
if the project is successful turtle products might be popularized to
an extent which might actually increase the pressure on wild
stocks, although Mariculture is shipping all of their products in
bond to prevent their being mixed with wild-turtle products and in
their advertising are emphasizing the superiority of the farmed
product over the natural product.
Another criticism that might be leveled is that the Mariculture
turtles are not allowed to graze on turtle-grass flats as they do in
the wild, but they are kept in circular cinder-block pools and are
fed largely on commercial trout chow and fish meal—i.e. they are
effectively practicing feed-lot monoculture, and their claim, im-
*Now known as Cayman Turtle Farm and based on largely German
financing; the eggs now come mostly from Surinam.
plicit in their name, of farming the seas is misleading. This effec

tively is raising the turtles to a higher trophic level than normal,
and will inevitably set a rather high minimum price for the turtles
when they reach marketable size after three years. Mariculture is
attempting to lessen the minimum economic price of the meat by
also selling numerous by-products, including the skins (which are
reported to be blemish-free and more valuable than the natural
product), the oil derived from the fat (which is used in the cos
metics industry and at the present time accounts for nearly 40% of
the firm’s cash intake), and the shells, which are either sold entire
to tourists, or the scutes are removed and used as imitation ‘tor
toiseshell.’ The shells of these captive-raised turtles differ slightly
from wild specimens, not only in having rather markedly scal
loped edges, but also in producing much thicker scutes, nearly as
thick as those of the hawksbill, probably because of the high pro
tein diet. The calipee too is an important secondary product, used
for enriching turtle soup. Moreover, those hatchlings which die
(and although the hatching rate of the eggs at Mariculture is very
high, the neonatal mortality is also very high) are embedded in
blocks of Perspex and sold as paperweights, mainly in Japan.
Despite these factors, however, it remains true that Maricul
ture is diminishing the total amount of protein available to the
human race, since the turtles eat more protein than they ultimate
ly produce. Moreover, the protein is taken from those who need it
most, and the final product is only available to those who need it
least—the gourmets and exotic food freaks of the rich countries.
Moreover, it has been seriously mooted by some outside conser
vationists that it is unlikely that Mariculture will ever be able to
maintain a breeding herd of sufficient size to produce all the eggs
they will need (about 100,000 per year at present), in which case
they would be forced to be dependent forever on eggs from the
wild. Whether or not this criticism is valid depends in large degree
on whether turtles can be induced (with hormonal injections, etc.)
to produce multiple egg clutches every year. If this proves impos
sible, the economics of maintaining several hundred adult-size
turtles in an artificial tank year after year may prove insuperable,
not only because of the tremendous amount of food that would be
required, but also because of the probability of severe and increas
ing problems in disposing of the excrement without polluting the
environment.
NON-FOOD PRODUCTS
The killing of sea turtles for non-food products—the hawksbill
for its shell and the ridleys for their skins—has caused a major
decline of both species, which is all the more regrettable because
both products are unnecessary luxury items for which perfectly
827
www.Ebook777.com
good cheap substitutes are available. The hawksbill industry is
centuries old; the species has survived to the present day mainly
because it is a diffuse rather than a colonial nester, and in the past
human populations in the tropics were sufficiently sparse for
many hawksbills to be able to nest in remote places without distur
bance. This former advantage is now a liability; few tropical
beaches are sufficiently remote nowadays for hawksbills to be
able to nest safely, and the absence of sites of aggregated nesting
(except for one or two islands in the Persian Gulf and Torres
Strait) makes enforcement of protection for nesting turtles more
or less impractical. Hawksbill products may no longer be
imported into the United States without a federal permit, and this
prohibition, which is now being enforced, may ultimately help to
reduce the demand for tortoiseshell.
The turtle skin industry has only existed for a few years, but in
that time literally millions of turtles have been killed for their
skins. The skin has an attractive ‘crazy paving’ pattern, easily dis
tinguished from crocodilian skin in which quadrangular scales are
arranged in linear rows. However, the skin comes in rather small
pieces, since only the skin of the front flippers and underside of the
neck is usable, and among wild turtles only the ridleys, which are
small even when adult, furnish ‘quality’ skins. Moreover turtle
skin is not very durable, being much thinner than cowhide, and the
process of flattening a turtle skin requires considerable stretching
in the elbow region, with consequent thin, weak areas. By far the
biggest trade in turtle skins is the export of Lepidochelys olivacea
skins from Mexico to Japan, though Australia and Costa Rica also
produce small quantities, and an appreciable market exists in the
United States and western Europe.
A highly regrettable aspect of the Mexican turtle industry is
that the price of skins is sufficiently high for it to be economically
unnecessary to utilize the flesh of the turtle as well, despite federal
laws requiring the use of all parts of the turtle. Some of the meat is
eaten locally or made into canned frankfurters, but typically
neither large markets nor canning facilities are locally available,
and the skinned turtles are thrown away—or, when government
fisheries agents are in the area, buried in huge pits in the ground.
Mexico is attempting to maintain its stock of ridleys by allowing
exploitation only under license and protecting turtles on the most
important nesting beaches (principally Playa la Escobilla, Oax
aca; Playon de Mismaloya, Jalisco; and Piedra de Tlacoyunque,
Guerrero). Nevertheless several factors reduce the effectiveness
of this conservation program. For one thing, huge numbers of tur
tles are killed illegally during the closed season, and their skins
are stored and released into legitimate outlets during the open sea
son. Moreover the program was initiated without any biological
828
data on the population dynamics of sea turtles, and there is no evi

dence that the numbers of hatchlings permitted to enter the sea
each year can offset the huge numbers of adults captured. Also col
lection of turtle eggs for food is almost universal on the beaches
where there is no conservation patrol, and the eggs, although a
totally illicit commodity throughout the country, are still openly
offered for sale even in inland cities; on one recent occasion a local
fisheries officer was found with 125,000 eggs, yet she escaped pros
ecution. However, the suggestion of the IUCN sea turtle committee
that the turtles could be maintained forever, and coastal Mexicans
kept happy as well, if the turtles themselves were protected and
exploitation of eggs only permitted, was not acceptable to the
authorities and to the industrial interests of Mexico. The sad
results of over-exploitation are already apparent; during the
twelve-month period August 1968 to August 1969, the nesting ag
gregation (or arribada) of ridleys at la Piedra de Tlacoyunque,
Guerrero, was reduced from an estimated 30,000 turtles to a few
hundred. But, to give credit where it is due, for several seasons
now perhaps the most effective turtle protection patrols in the
world have been conducted to save the last few Kemp’s ridleys
(Lepidochelys kempi) on the nesting beach in Tamaulipas.
The Atlantic shrimping industry is the direct cause of numer
ous sea turtle deaths; turtles are caught in drag nets and often are
held under for so long that they drown. If they are alive when
hauled out they may still be killed by crewmen, who dislike turtles
because they tear their nets. Most of the at-sea recoveries of
ridleys tagged in Surinam were made in this way, and on rare
occasions the turtles are released alive after the tag number has
been recorded. St. Augustine-based shrimp fleets frequently catch
loggerheads off the coast of Florida, and if these turtles are still
alive when hauled aboard they are often submitted to frightful
mutilations before being thrown back in the sea. It is not unusual to
find drowned loggerheads, whose front flippers have been com
pletely severed, thrown up on Florida beaches.
SCIENTIFIC COLLECTION
Appreciable numbers of turtles are required for scientific
study. There are now many dozens of museums throughout the
world which would theoretically like to have a series of preserved
specimens of each turtle species, though in many cases a museum
does emphasize one particular area in its collections, with only
scant representation of species from outside this area. Turtles are
also used in very large quantities for anatomical dissection and for
neurological and circulatory demonstrations in undergraduate
classrooms in American universities. The difference between
829
www.Ebook777.com
these two types of exploitation is that in the former case the pres
sure is on the rare species, while in the latter case the cheapest and
most easily obtainable turtles are used (usually Chrysemys picta
bellii and Pseudemys scripta elegans). There are as yet relatively
few instances where the demands of museums have had noticeable
effects on turtle populations, but there have been some cases
where museum collectors have taken far more of certain species
with very limited ranges than were required for adequate charac
terization and description of the form; here we may mention the 86
(mostly female) Duncan Island tortoises (Geochelone elephan-
topus ephippium) collected by the California Academy of Sciences
expedition in 1906, and the huge series of certain highly restricted,
though locally common, Graptemys species collected by the
Tulane University crews in the 1950’s; hundreds of individuals
were also killed for dissection and not even retained for the Tulane
Museum collection. But happily the old maxim ‘kill it first, study it
later (if at all)’ is at last losing favor, and biologists usually think
twice nowadays before slaughtering large series of adult turtles,
perhaps several decades old, just to see what they had for their
last meal.
PET INDUSTRY
The demand for turtles as pets and zoo animals increases from
year to year, but as yet few species appear to have been seriously
depleted by this demand. The bulk of the pet industry is still based
on hatchling red-eared turtles (Pseudemys scripta elegans), and
since it would be extremely difficult to catch sufficient hatchlings
in the wild to satisfy the demand, these turtles are raised on large-
scale farms in Louisiana and Mississippi. High densities of adult
turtles are maintained in outdoor pools with banks suitable for
nesting, and large crops of hatchling turtles are produced annual
ly. There is thus no objection on strictly conservation grounds to
the hatchling turtle industry in this country, though the conditions
under which the baby turtles are maintained before (and after)
sale may well give rise to complaints based on humanitarian con
siderations. Also, the practice of releasing deformed hatchlings as
a ‘conservation measure’ is dubious, to put it mildly.
The recent Salmonella scare has apparently put a serious re
straint on commercial pet turtle farms, and it is possible that
before long they will be put out of business altogether. The indus
try has responded by recommending that if baby turtles are kept
without water except when actually fed, there is no danger of them
harboring Salmonella. Needless to say, this weird and purely self-
serving recommendation is totally without merit, as the dry con
ditions are more likely to kill the turtle than the Salmonella, and no
one has yet succeeded in raising an aquatic turtle in a dry tank.
The rising number of serious turtle hobbyists in the United

States usually consider a wide variety of species to be the most im
portant criterion of a good turtle collection, and the demand is thus
spread over all species without excessive pressure on any one;
moreover the high prices asked by most exotic turtle dealers now
adays help to ensure that buyers will only be those who have a seri
ous and sustained interest. One particular case, however, where
the demand for exhibition or pet animals may have an undesirable
result is the case of the Galapagos tortoises.
There are about 10,000 to 15,000 Galapagos tortoises left in the
wild, yet many small zoos in the United States have several of
these animals even if they have few other reptiles. Wealthy private
collectors too are willing to spend large sums on Galapagos tor
toises, and the result is that tortoises are still being illegally ex
ported both from the islands themselves and from Ecuador, where
apparently there are a number of tortoises that were brought from
the islands before their export was banned. Very few zoos and no
private collectors have ever bred Galapagos tortoises, so a group
of people who often call themselves conservationists, and who do
have a genuine regard for the tortoises in their collections, are in
fact providing one more obstacle to the Galapagos tortoises’ strug
gle for survival. The answer to the problem is probably to register
all Galapagos tortoises presently in the country, perhaps marking
each in a unique way or recording individual peculiarities of each.
After an amnesty period, any tortoises found unmarked or unre
corded should be seized by a conservation agency empowered by
the federal government and returned to the islands. Captive-bred
tortoises could be recorded as they are hatched. This system would
probably work, as there must be few owners of Galapagos tor
toises who would be content to keep their animals in complete
secrecy, since the urge to show off one’s rare specimens is usually
very strong.
The idle killing of turtles by vandals is a problem in both the
United States and Australia, where there appear to be as many
people who will swerve to hit a turtle crossing the highway as there
are who will swerve to avoid one; there are even those who will
drive their car through a group of migrating turtles, zig-zagging
about to see how many they can hit. Many turtles are also killed on
highways by people with poor driving habits, though of no particu
lar bad intention, who drive so close behind the car in front that
they do not see a turtle on the highway until it is too late.
Even in National Parks in the United States tortoises are used
for target practice, and in the south rednecks sometimes amuse
themselves shooting at basking cooters, or even at the heads of
swimming or floating turtles. There is even a rather attractive
book on the market entitled The Pest and Varmint Hunter’s Bible
sai
www.Ebook777.com
which gives instructions on shooting the heads off snapping turtles.
The habit of shooting turtles stems from several causes; lack of
education or environmental respect, easy availability of firearms,
and simple boredom. The urge to kill for recreation does appear to
be confined to those who lead humdrum or unsatisfying lives. No
easy way of eliminating this type of vandalism springs to mind;
the ultimate solution involves far-reaching cultural and social im
provements that cannot be organized overnight. A good start
might be to declare the practice illegal, and to arrange temporary
employment in abattoirs for offenders.
Habitat upset and destruction have already depleted popula
tions of several turtle species and undoubtedly will jeopardize
many more in the future. Upset of the Galapagos ecosystems, and
its effect on the tortoise populations, was discussed earlier. In the
United States increasing urbanization has reduced the populations
of many species; this effect has been most marked on the southern
California pond turtle (Clemmys marmorata pallida), the red-
bellied turtle (Pseudemys rubriventris), and the bog turtle (Clem
mys muhlenbergi). The range of the bog turtle has been steadily
shrinking this century, and will do so further as more of its special
ized habitat is drained and reclaimed. The species can be bred in
captivity, but it is also necessary for wild areas to be purchased
and set aside as reserves before they are destroyed.
In some parts of the world sea turtles face habitat destruction
on the nesting beaches. Some Far Eastern beaches are so
cluttered with industrial debris that there is no clear nesting space
left. In the Caribbean area the spread of the coconut palm has in
some places had a bad effect on turtle beaches, since the root sys
tems of these trees spread throughout the sand above the high tide
mark, making it difficult or impossible for a turtle to nest. On the
Atlantic coast of Florida beach development and highways some
times produce such intensive illumination behind the beach that
newly emerged hatchling loggerheads are attracted away from
the sea and lured to their deaths beneath the wheels of cars on the
road. This is a genuinely accidental result of urbanization and
often occurs when the local people have feelings of nothing but
good will toward sea turtles. The best cure would probably be to
eliminate unnecessary illumination behind the beach, but if this
proves impossible, it should be quite easy to construct a continuous
low wall or curb at the back of the beach—it need be only a few
inches high. The baby turtles will be prevented from crossing the
highway by this device, and at dawn the rising sun should give
them a strong enough cue to head for the sea.
832
Chapter 12
TURTLES IN CAPTIVITY
METHODS OF OBTAINING TURTLES
The average turtle keeper obtains his specimens from a pet
shop or mailorder dealer. In recent years a tremendous variety of
species has become available (though recent legislation is likely
soon to restrict this number considerably), and some forms, in
cluding Podocnemis unifilis and Pseudemys scripta callirostris,
became large-scale items of commerce even before they were rep
resented in most museums. Most mailorder dealerships guarantee
live arrival and will accept return of animals with defects not men
tioned on the price list, so there is no reason to hesitate in buying
from such sources—though of course it is more satisfactory to visit
the establishment if this is possible and make a personal selection.
In choosing a turtle from a pet shop, the most important criteria
are clear eyes and nose and heavy body; a turtle whose eyes
appear gummed up or watery, which shows any signs of bubbling
at the nose, or which feels light and floats high in water (or even
worse floats on one side) is not in good health and probably will not
survive. With aquatic species it is also essential to check that the
turtle does not suffer from skin fungus or shell rot of any kind—
especially a rather insidious kind of shell rot that erodes the bone
while leaving the superficial scute intact. A healthy turtle should
be quick in its responses and should struggle or withdraw when
picked up, though such responses will necessarily be slower if the
ambient temperature is low.
If the opportunity presents itself, the turtle hobbyist usually
likes to capture at least some of his specimens himself. The key to
successful collection of terrestrial turtles is to recognize likely
habitat and to learn when the turtles are likely to be abroad and
where they are likely to hide when they are not; apart from this,
perhaps the best method, especially if one spends plenty of time
driving, is to pick up those turtles found crossing highways. This
method has the advantage of salving one’s conscience, since one is
probably saving the life of the turtle instead of withdrawing it from
the security of its natural habitat. It is also a humanitarian gesture
to pick up those turtles on highways which one does not need, and
to set them down on the side of the road for which they heading
R33
www.Ebook777.com
when first seen. At certain times of the year, especially during or
after heavy rain following a summer drought period, large num
bers of both aquatic and terrestrial turtles may be found crossing
highways in this way. Many museum collectors nowadays pick up
so-called DOR (“dead on road”) specimens; having good locality
data, these are valuable and enable good series to be built up with
out the sacrifice of live specimens. Those which are too rotten to
lend themselves to preservation can usually be made into
skeletons by placing the specimen in warm water for a few days.
Certain rural people in the southern United States are highly
expert at extracting gopher tortoises from their burrows by means
of wire ropes with hooks on the ends, but this method is likely to
damage the tortoise and is not recommended to the turtle hobby
ist, for whom gopher tortoises are not recommended anyway since
they rarely live long in captivity.
Aquatic turtles may be caught in many different ways. The
most straight-forward method is simply to wade around the edges
of a pond or lake and grab any turtles which are seen; the usual
reaction of a turtle is to swim a short distance and then seek refuge
on the bottom, and if it is kept in sight it can often be caught.
Another useful method, applicable in clear waters such as the
springs of north and central Florida, is to don face mask and flip
pers and to catch the turtles by swimming after them. With consid
erable skill, the method can also be adapted for capture of even
very shy turtles, such as Graptemys, in the less clear waters of the
rivers draining into the northern Gulf of Mexico. Moreover most
aquatic turtles spend long hours basking in the sun, usually on
half-submerged logs and snags but sometimes also on river banks.
Such turtles are usually very wary and difficult to capture, but
with skill may sometimes be caught. When a basking turtle is so
situated that one can approach it closely while keeping completely
out of sight, it may often be caught with a sudden final grab. There
is no need for silence during such a stealthy approach, since tur
tles, although not deaf, usually show amazingly little response to
sound. Their eyesight, however, is very good, and I have often seen
basking turtles drop back into the water while I was still hundreds
of yards away. Shy river turtles may sometimes be captured by a
floating, rather than swimming, approach. If one enters the water
half a mile or more upstream (or round the nearest upstream bend
in a winding river), and floats down in an inflated inner tube, pre
ferably with some suitable camouflage such as a mass of leafy
vegetation, one can often approach so close to basking turtles that
they may be caught in a dip net as they drop off their basking logs.
Many carnivorous turtles, especially mud and musk turtles,
can be caught on hook and line. To avoid damage it is better to use
a barbless hook. Fishermen frequently catch such turtles uninten
834
tionally, and usually to their intense irritation. Alligator snapping

turtles may be caught at night, when the weather is reasonably
warm, by suspending bait (fish or liver) in the water from over
hanging branches. The bait site can be re-visited during the night,
and any turtles seen (with the aid of a bright light) can be brought
to the surface by a swimmer or with a blunt gaff.
Turtles can be trapped with a variety of devices. It is possible
to make traps in the form of wire cages which are attached to
basking platforms; when the basking turtles are disturbed, they
will dive off and hopefully land in the cages. It is better to attach
the trap to a favorite basking log rather than to plan an artificial
basking structure, since turtles are usually very wary of unfamil
iar basking surfaces and tend to avoid them. However, a favorite
trap in former times was a barrel with a plank nailed across the
open top and sunk flush with the surface of the water. Turtles
diving off the plank were trapped in the barrel.
Of more general application are baited traps which trap tur
tles which are searching for food. A typical device, useful especial
ly for snapping turtles, consists of a large chicken wire cage with a
lockable door and a funnel-shaped opening into which a turtle can
swim. The inner end of the funnel can be equipped with a door that
can be pushed open only from outside; but even if no such door is
provided, very few turtles will escape. Such a trap, which may be
cylindrical or cuboidal, is staked in shallow water with one third or
less of its volume exposed above the surface; a totally submerged
trap will drown turtles. Many types of bait may be used; water
melon rind is said to be particularly effective in trapping soft-
shells, but sardines are a more useful general-purpose bait. Tur
tles may also be caught in seine nets or in trammel nets, which
have floats attached to the upper margin and are designed to reach
close to the bottom of the water. These long nets are stretched be
tween two waders or two small boats and allow a large area to be
swept and many turtles to be caught.
Finally, a favorite method of obtaining turtles in tropical coun
tries is to visit market places where the animals are sold for food.
For example, in the states of Veracruz and Tabasco in southern
Mexico it is possible to purchase snapping turtles, Dermatemys,
Staurotypus, Pseudemys, Kinosternon (several species), and even
the little Claudius, which is almost never seen for sale in the
United States. In Guatemala City I have seen large numbers of the
small, beautifully marked Rhinoclemys pulcherrima offered for
sale as food, as well as Staurotypus salvini, Kinosternon s. cruenta-
tum, and Pseudemys scripta grayi. In Surinam tortoises (Geoche■
lone carbonaria and denticulata) are favorite items of food, espe
cially among the Chinese community, and since they are invari
ably kept alive until eaten it is sometimes possible to buy them. In
ooer
www.Ebook777.com
Haiti, large numbers of Pseudemys decorata are sold in the mar
ket in Port-au-Prince; even hatchlings are sold, these apparently
being raised at home on scraps before being eaten. In the market
in Guayaquil, Ecuador, one can usually buy snapping turtles (Che
lydra serpentina acutirostris). Also, in many tropical or unfamil
iar areas the turtle hobbyist can often obtain a good cross-section
of local species by offering a small reward to local people for each
turtle found.
LEGISLATION
In contrast to the old free-and-easy days, the turtle keeper
nowadays finds his hobby hedged around with many legal restric
tions. This book obviously cannot undertake to be a complete ref
erence on existing regulations, but some idea of their scope can be
given.
The United States Department of the Interior has now
declared a number of animals, including several turtle species,
‘endangered species,’ which are effectively off-limits to amateur
hobbyists, since importation permits are only granted to institu
tions and individuals who can demonstrate that the importation of
the turtles they want will not have adverse effects on wild popula
tions of the species and that their possession of the turtles will
materially contribute to the survival outlook for the species. This
list is constantly being updated; as of June 4,1973, the list includes
all races of Galapagos tortoises, Geochelone elephantopus; the
radiated tortoise, Geochelone radiata; the western Australian
Pseudemydura umbrina ', the South American Podocnemis expan-
sa and unifilis; the sea turtles Lepidochelys kempi, Eretmochelys
imbricata, and Dermochelys coriacea ', the aquatic box turtle Ter-
rapene coahuila', and the Southeast Asian Batagur baska. At the
present time a species has to be ‘threatened throughout its natural
range’ to be included in the list; but a modification is now being
considered to include threatened populations of species that are
safe in other parts of their range. Legislation (opposed by special
interest groups) is now pending to place the green turtle (Chelonia
mydas) and the loggerhead (Caretta caretta) on the list. Some of
the above species were placed on the endangered list because their
ranges are so restricted that any collection of specimens is con
sidered deleterious. Others, including the sea turtles and the two
Podocnemis species, are important and over-exploited articles of
commerce, not yet on the point of complete extinction, but declin
ing and therefore in need of protection. Ownership of endangered
species is not prohibited if the specimens were already in the coun
try when the ban was established. Needless to say, private owner
ship of species on the list is not encouraged, and some zoos are
836
This hatchling Geochelone e. hoodensis shows there is still hope for several
Galapagos tortoise populations.
transferring endangered species (such as Galapagos tortoises) to

other institutions more favorably situated for breeding.
Recent legislation further restricts importation of wildlife,
including many turtles. This legislation includes so-called “Appen
dix I” animals (endangered species) and “Appendix II” animals
which may be imported if a license is obtained from the country of
origin. In the United States the Lacey Act also applies—if an ani
mal is protected in its country of origin, it may not be imported
into the U.S. without a special permit. Appendix I currently in
cludes the following turtle species: Geoclemmys hamiltoni, Mela-
nochelys tricarinata, Kachuga tecta tecta, Morenia ocellata, Bata-
gur baska, Terrapene coahuila, Geochelone elephantopus, Geo
chelone radiata, Geochelone yniphora, Psammobates geometri-
cus, Lissemys punctata punctata, Trionyx ater, T. nigricans, T.
hurum, T. gangeticus, Pseudemydura umbrina, and all sea turtles
except for Australian Chelonia populations. It will be noticed that,
apart from the obvious candidates, a number of species from the
Indian region are included which are not normally considered rare
or threatened. Their inclusion apparently results from a misunder
standing of the real nature of the list by the Indian authorities when
they were asked to submit names for inclusion.
The species of turtle currently to be found in Appendix II
S37
www.Ebook777.com
include Clemmys muhlenbergi, all Podocnemis species, and all
tortoises and sea turtles not Included in Appendix I. These regula
tions may seem irksome to the prospective turtle keeper, but it
should be borne in mind that they exist for the benefit of the ani
mals concerned and were in part brought about because of abuses
by irresponsible dealers who imported enormous numbers of cer
tain vulnerable and attractive species in the hope that at least a
few would survive long enough to be sold. Further legislation cur
rently pending in the United States would ban all importation of
wildlife with the exception of certain domestic or otherwise
exempt species unless a permit was first obtained. The thinking
behind this legislation is only partially oriented toward conserva
tion of the species concerned; it also takes note of the danger that
certain exotic species are liable to become established in the
United States, a danger especially great in such vulnerable areas
as Hawaii and South Florida. Permits are likely to be available
both to professionals and to those amateurs who are able to demon
strate that they have made suitable provision for housing their
specimens. It is anticipated that this legislation will not complete
ly eliminate animal dealers, since they will still be able to sell
native U.S. species and also will be able to act as agents for the im
portation of specimens for permit-holding buyers. It is also likely
that provision will be made to allow importation of the mixed and
unpredictable collections that field workers connected with
museums, zoos, and certain private collections may assemble.
In most parts of the United States it is legal to capture turtles
for non-commercial purposes, but certain states have passed re
strictions on certain species. In Florida it is not legal to export out
of the state gopher tortoises, Barbour’s map turtles, or alligator
snapping turtles, to buy or sell them, or to own more than one spec
imen of each without obtaining a permit from the Game and Fish
Commission. Texas completely protects the Texas tortoise, and the
desert tortoise is completely protected in California and Nevada.
New York State prohibits capture of box or wood turtles. Several
other states have regulations against the shooting of turtles and re
quire permits for the hunting or collection of turtles for commer
cial purposes. The bog turtle is protected in New York, New
Jersey, and Pennsylvania.
CARE OF TORTOISES
The easiest way to maintain a tortoise in captivity is to release
it in an enclosed garden or yard in an area whose climate offers a
reasonable facsimile of that to which the animal would be subjec
ted in its natural habitat. Under such conditions tortoises can live
838
for many years, obtaining their own food by browsing on whatever

vegetation is available. However, tortoises tend to be rather sen
sitive to suboptimal temperature conditions or perhaps even more
to unnaturally humid situations, and for this reason many turtle
keepers find it much harder to keep tortoises alive than aquatic
species. Nevertheless, many hobbyists do manage to raise and
even breed tortoises, and some general rules can be drawn up
from those who have been successful.
In nature tortoises are typically almost entirely herbivorous.
Some are quite content to eat nothing but grass, but most tortoises
eat a wide variety of green vegetation, with certain distinct prefer
ences (most tortoises are very fond of lettuce, even though this is
not a very useful food from a nutritional point of view). Tortoises
often show great fondness for flowers, and many species will sub
sist happily on fruit. Many tortoises show a preference for red or
yellow colored food. However, it seems quite probable that the her
bivorous diet typical of most tortoises is not so much a preference
as a necessity, since no tortoise is fast enough to catch live prey. In
the wild state it is probable that carrion is eaten whenever the op
portunity presents itself, and many tortoise keepers have found
that their pets put on wonders of growth when fed on commercial
canned dog food. However, they may need to be weaned on to this
gradually as they may not initially even realize that the dog food is
edible. Big tortoises have great appetites, and a good and also eco
nomical diet can be worked out with canned dog food and slightly
over-ripe fruit or lettuce outer leaves which may be obtained free of
charge from a supermarket produce department. Since tortoises
have such a high proportion of bone in their bodies, it is a good idea
to sprinkle the food with a commercial bone meal and multivita
min supplement.
Tortoises differ in their water requirements. Giant tortoises
from the Galapagos Islands or Aldabra spend much of their days
in the wild half-submerged in fresh water, and for these animals it
is a good idea to make a warm-water pool perpetually available for
both drinking and bathing. For most other tortoises standing water
is only intermittently available, and consequently they will not
suffer if kept dry for long periods; indeed some tortoises will over
drink if water is kept constantly available, and they may become
waterlogged and eventually die. For most tortoises, therefore, it is
quite sufficient to provide them once a week with a few hours’
access to water in which they can partially submerge themselves.
For those species from relatively humid areas, such as Geoche-
lone denticulata, G. emys and G. impressa, it will do no harm to
have water constantly available. On the other hand, tortoises from
desert areas, particularly the little South African tortoises of the
genera Homopus and Psammobates, require very hot, dry condi-
839
www.Ebook777.com
tions if health is to be maintained. The temperature may be
allowed to drop at night, but the combination of low temperature
with high humidity is usually rapidly fatal.
In most parts of the United States it is probably best to keep
tortoises outside during the summer months. If one’s yard is not
fenced (and fencing needs to be very secure to prevent small tor
toises from escaping), the animals can be enclosed with a metal
swimming pool siding, installed in a shallow trench with the dirt
packed back in. Alternatively, brick or block walls can be
constructed. It is important that tortoises always have a retreat
from the sun, provided either by dense vegetation or by an artifi
cial hutch. Tortoises may also be kept indoors, and tropical tor
toises have to be kept indoors in winter. A dry, draft-free heated
greenhouse is probably the ideal structure for this purpose, but if
this is not available tortoises may be kept in a dry aquarium with a
soil substrate, or in a plywood box, according to their size. A sun
lamp should be provided for an hour or more a day, and if temper
atures are likely to drop low at night an infra-red heat lamp should
be used. Some tortoise keepers have successfully kept their pets in
walk-in closets with the floor covered with plastic and several
layers of newspaper.
Temperate zone tortoises may be allowed to hibernate in win
ter, if they appear to be in good condition and with adequate fat re
serves in the fall. Ideal hibernation temperatures are a few
degrees above freezing, so in the north an unheated cellar provides
a good hibernaculum. Drafts and rats should be rigorously exclu
ded, and an insulating medium such as straw or tom-up news
papers should be provided. Temperatures below freezing even for
short periods are liable to be fatal, while if the temperature rises
too much the tortoises may wake up sufficiently to start metaboliz
ing rapidly but not enough to start feeding. Under such conditions
they may lose a lot of weight and may not recover.
CARE OF AQUATIC TURTLES

A wide variety of containers can be used for housing aquatic
turtles. For small specimens, a large aquarium equipped with fil
ter and re-circulating water may be used. For larger ones, a good
and very economical tank can be made from the inner liner of an
old refrigerator. These liners usually have a variable number of
bolt holes which can be filled with silicone cement. They make a
much more decorative tank if the front is cut out with a blowtorch
and a piece of glass fixed in, again with silicone aquarium cement.
Still larger turtles may be kept in cattle watering tanks, or better
in permanent cement pools. A very attractive display can be made
840
with a built-in pool in a greenhouse, and with such an arrangement

even tropical turtles can be kept year-round in all climates.
Turtles from a climate similar to that prevailing locally can be out
of doors year-round in a deep garden pool, at the bottom of which
they can hibernate during the cold months. However, the pool
should be at least two feet deep at the deepest point and with a suit
able substrate of mud or debris to protect the hibernating turtles
from freezing.
Most turtle keepers find that tank decoration is best
minimized, except in the case of very small or hatchling turtles in
a large tank. Turtles tend to uproot or eat aquatic vegetation, and
gravel or sand on the bottom of a tank can become very hard to
keep clean, so the ideal arrangement is a tank devoid of organic
decoration which can be completely drained with a minimum of
trouble, rinsed, and re-filled. Active turtles in moderate or high
concentration will foul their water every few days, so provision for
easy and rapid changing of the water is very important.
Nearly all aquatic turtles appreciate having an emergent plat
form for basking, and basking in direct sunlight (or an artificial
source rich in ultraviolet) is important to their health. The basking
platform may be made from natural wood or rocks, or an equally
functional though less decorative surface can be made from a sus
pended piece of board or plastic, or from folded wire mesh, the lat
ter having the additional advantage in a small tank of allowing the
turtles room to swim or hide under it. It is desirable also to have a
land surface available that is shaded from direct sunlight, though
turtles can certainly survive without such a provision. Some highly
aquatic species, such as the alligator snapping turtle and some
kinosternids and trionychids, rarely if ever bask, and will do per
fectly without a basking platform. Needless to say, a more com
plex arrangement, with an extensive land area of sand or soil, is
essential if it is the owner’s intention to try and breed aquatic tur
tles of any kind. If this is not available, they will either not lay eggs
at all or else will lay occasional over-mature and hypercalcified
eggs in the water.
As far as temperature maintenance is concerned, many
tropical species can survive surprisingly low temperatures (50° F.
or even below) without ill effects, though of course they will not
feed at such temperatures. Others, such as certain Podocnemis
species, are unable to survive cold temperatures even for a short
time. The turtle keeper should play safe with all his species which
are not exposed to cold conditions in the wild and bring them in
doors or heat their water whenever the temperature drops below
about 55°-60° F. Turtles are much more tolerant of thermal shock
than fishes, but nevertheless excessively rapid thermal changes
should be avoided.
841
www.Ebook777.com
Certain aquatic turtles, particularly half-grown and adult
Pseudemys, Dermatemys, and Podocnemis, enjoy a largely vege
tarian diet and may be fed on a variety of fruit and vegetables, es
pecially lettuce and banana. Many other species are purely carniv
orous and flourish best on a somewhat mixed diet. A good standby
is raw fish—whiting or smelt. Mullet is also nutritious and cheap,
but is so oily and scaly that tanks can be difficult to clean up after
wards. Purina trout chow is a nutritious and very cheap food for
turtles, and is also relatively non-polluting of tank water, but tur
tles usually take time even to recognize trout chow as edible. In
addition to these foods, it has also been recommended that turtles
be fed a composite gel-type food about once a week to ensure that
supplements are provided and are bound together so they are not
lost before ingestion. The gel recipe given by John Hoke (Turtles
and Their Care) is as follows:
2 oz. fish (frozen smelt or any fresh fish)
8 oz. boiling water
3 oz. trout chow, ground up into a powder
2 oz. cod liver oil
1.5 oz. unflavored gelatin
1.5 oz. bone meal
1 teaspoon dibasic calcium phosphate
20 drops of red food coloring
Mince the fish into very small pieces. A blender can be used
for this. Add the cod liver oil to the fish and set aside. Dissolve un
flavored gelatin in the boiling water, stirring thoroughly. Next,
add the minced fish and oil. Then pour in the ground trout chow,
along with the bone meal and dibasic calcium phosphate. Mix well,
adding twenty drops of food coloring. Pour the mix into a shallow
pie tin or plastic ice-cube tray and put it in the freezer. Gel should
be cut into small cubes or strips to feed to the turtles. Give as much
as they will eat at one time. Leftover visible scraps should be
netted out of the tank so that they will not have to be processed by
the filtration system. Always return uneaten gel to the freezer
quickly, covering it with plastic wrap so that it will not dry out.
BREEDING
Turtles may lay eggs in captivity, either singly on the surface
of the ground or as a whole clutch properly buried as in the wild. If
eggs are properly buried and the climate is suitably warm, they
may be left in situ to hatch; the nest area should be surrounded by
hardware cloth to prevent escape and loss of the little turtles as
they emerge. If conditions are less ideal, the eggs may be transfer
red to an incubator. The eggs should be moved as soon as possible
after laying, with the minimum of mechanical disturbance, and
842
Hatching Testudo marginata.

Note the “egg tooth” on the
snout. Photo by Othmar Stem
mier.
their original orientation should be preserved throughout the

transfer. Movement of eggs more than a day or so after laying
may kill them, though after a few months, when the embryo is al
ready well-developed, they may again be moved with care. Tor
toise eggs should be kept dry and warm — ideally at about 80°, with
a nightly drop to not less than about 75°. Eggs of aquatic turtles
should be kept a little more moist; a good way to keep them at a
suitable humidity is to blow up a plastic bag and place the eggs in
side, then tie off the mouth of the bag. In this way, humidity from
one’s breath will be maintained throughout incubation. Hatching
may take only ten weeks or so in some cases (eight weeks or less
for sea turtle eggs), but in other cases five or six months may
elapse before hatchlings emerge, and unhatched baby turtles are
also capable of over-wintering in the egg. Many turtle owners have
been tempted to give up and open eggs that had been incubating
for several months, only to find that if they had waited a few more
days or weeks the eggs would have hatched. Consequently only
those eggs which have definitely spoiled should be rejected.
DISEASES OF TURTLES
Captive turtles are liable to a considerable number of diseases
which are rarely if ever found in wild-living individuals. One of the
commonest is pneumonia, which tortoises are particularly prone
to catch if they are kept under damp or cold conditions. It is mani
fested by the animal refusing to feed, bubbling at the nose, and
often producing a rasping noise as it breathes. It is usually fatal if
843
www.Ebook777.com
untreated. The best chance of curing the condition is oral adminis
tration of a mixture of antibiotics and vitamins. Four tablets of
Aureomycin mixed with four eye-dropperfuls of liquid vitamins
may be mixed with a little water and dispensed by means of an
eye-dropper several times a day. If the tortoise is not too far gone,
this should clear up the condition within a week. An essential part
of the cure is to provide the tortoise with sufficiently sunny, warm,
dry quarters to prevent the condition which led to the disease in the
first place. As with most turtle diseases, prevention is easier than
cure.
Aquatic turtles often suffer from whitish patches of fungus on
the skin; with sea turtles, Pseudomonas is the usual causative or
ganism. These fungoid areas may be treated with common salt, or
better with iodine, Gentian violet, malachite green, or merthiolate
painted on the affected area, or the turtle may be placed in dilute
(strongly pink) potassium permanganate solution. A thick green
ish mixture known as Coppertox may also be used for skin fungus.
Various deficiency diseases are common among captive tur
tles. Hatchling turtles raised in captivity often acquire deformed,
soft shells though they may continue to feed well and grow. This
condition is caused by a vitamin D deficiency and is best cured by
Kyphotic specimen of Trionyx This hump-backed condition is moderately

common in soft-shells.
844
regular vitamin D supplements as well as a calcium phosphate

(bone meal) food supplement and exposure to several hours of sun
light per day. It is of course essential to provide a retreat from ex
cessive sunlight. Both terrestrial and aquatic turtles can profit
from free access to a limestone block; gnawing on this prevents
beak overgrowth and provides valuable supplementary calcium.
Captive turtles sometimes suffer from swollen or discharging
eyes; in severe cases the eye may become completely closed and
clogged with cheese-like material. This is caused by a deficiency
of vitamin A. A curative dose consists of 5000 International Units of
vitamin A per pound of body weight, injected intramuscularly.
Vitamin A may also be given orally, together with sugar water and
cod liver oil. A warm environment also assists in the cure. Hatch
ling turtles with severe hypovitaminosis A may be so weakened
that a cure is difficult; however, larger turtles, even those whose
eyes appear to be so infected that they could never be cleared, may
recover completely and regain normal eyesight. Other symptoms
of hypovitaminosis A are a general breakdown of integumentary
systems, with excessive skin peeling, as well as loss of the lining of
the intestinal and urogenital tracts. For local eye relief in cases of
this disease, boric acid or a weak warm solution of common salt
may be applied. Eye infections of various kinds may also be
reated with antibiotic ointment (Neomycin, Polymyxin B)
applied 4 times daily.
A deficiency of vitamin B12 may manifest itself by paralysis of
the legs. The cure is to present this vitamin orally, plus supple
mentary calcium and trace amounts of copper sulphate.
Shell rot is particularly common in chelid side-necks and in
soft-shells and certain emydids and kinostemids. The cause is not
completely understood, but it is thought to be associated with alka
line water. John Mehrtens suggests that the problem can be avoid
ed by addition of sufficient monobasic sodium phosphate to bring
the water to a pH of 7.0, or better to 6.6 or 6.8. Shell rot may mani
fest itself by formation of lesions in the plastral bone which ulti
mately may perforate completely, the bone degenerating to a
cheese-like material. The overlying scute, however, may remain
completely intact, but nevertheless the infected area is usually
visible through the scute. The rotten material and overlying part
of the scute should be completely excised using a scalpel or razor
blade and tweezers, the area cleaned and dried thoroughly and
painted with Gentian violet, or else packed with Neomycin/Corti-
sone ointment and covered with a layer of zinc oxide paste. Once
the causative organisms have been destroyed, the bone can regen
erate quite quickly.
Localized shell abrasion is usually self-curing once the cause
of the abrasion has been removed. In many cases the external
845
www.Ebook777.com
layer of bone will die when the overlying scute has been removed
or damaged; however, this dead bone will remain in place a long
time as a protection for the delicate live bone underneath, which
will gradually secrete a new horny layer. When the dead bone is
peeled away, a greying or silvery color indicates that scute regen
eration is proceeding properly. However, if infected material is
present, it should be removed and the area disinfected.
Soft-shell turtles may suffer from the disease of “red-leg,”
which is more common in amphibians and is characterized by a
breakdown in the superficial capillaries giving a blotchy, pinkish
appearance to the skin. The cure is to immerse the turtle in a
1:10,000 acriflavine solution for several hours daily. Severely frac
tured shells, such as may be suffered by a glancing encounter with
an automobile, should be treated with boric acid or Canadian bal
sam and taped up with waterproof electrician’s tape. This may be
left on for weeks or even months if necessary.
Most of the various parasites which have been found in turtles
are not known to cause severe harm to the animal, and most of the
internal ones may be very hard to detect. However, if the turtle
passes visible worms, the best treatment is to add piperazine to the
drinking water. Nematode infections may be treated by adminis
tration of equal quantities of calomel and santonin (about one
gram of each for a medium sized turtle), followed 24 hours later by
a teaspoon of castor oil.
Visible external parasites, such as ticks, should be painted
with paraffin or methylated spirit to weaken their grip, then re
moved with tweezers. Ticks are more common on wild than on
captive tortoises.
Subcutaneous lumps or tumors are sometimes caused by the
presence of pockets of maggots of the bot fly. These lumps should
be opened with a scalpel and the contents removed with forceps.
Captive turtles may suffer from hard, swollen lumps under the
skin of the limbs and neck. When these are close to the surface they
may often be squeezed out; in other cases it may be necessary to
make an incision first. Healing usually takes place once such
lumps have been removed, though the area should be treated with
Gentian violet to assist healing when the lumps are particularly
large. Small skin wounds or incisions may also be treated with
mercurochrome.
A turtle that has trouble sinking in water, or which floats with
one side much higher than the other, and also suffers from swollen
feet, is probably suffering from a Proteus infection. This is best
treated with Neomycin or dihydrostreptomycin; it is a serious con
dition, and untreated turtles usually die.
The turtle disease that has attracted the most attention in re
cent years is salmonellosis, since this disease can be passed on to
846
humans. Salmonella, the organism responsible for salmonellosis,

is a genus of bacteria of which about 1400 “species” (serotypes)
are known. Closely related is the genus Arizona, distinguished by
its inability to ferment lactose, of which about 400 serotypes have
been identified. Salmonella is a very widely distributed organism
and certainly is by no means confined to turtles. Infection can be
serious in humans, particularly very young or very old people; it
manifests itself by fever, stomach and intestinal upsets, nausea,
etc.
Salmonella is present in about 80% of samples of supermarket
chicken and can even be cultured from about 13% of a random
sample of coins and banknotes. It has also been identified in imita
tion ice cream, commercially baked meringue pies, nonfat dried
milk, eggs, fish, pork, beef, potato salad, lemon tarts, spaghetti,
cheese, candy, and lipstick. The organism multiplies either in the
gall bladder or the kidney pelvis of infected host animals; the for
mer is the more common location. Turtles, of course, have to be
exposed to Salmonella before they can carry it, and herein lies the
cure. If turtles are maintained in clean conditions and not fed on
raw offal and other fertile sources of Salmonella, they are unlikely
to acquire the organism or transmit it to humans. I might add
parenthetically that I have kept dozens of turtles for many years
and have fed them on a wide variety of foods with no attempt to
check for possible Salmonella introduction, and have never suf
fered from any symptoms of salmonellosis. However, it is possible
that children who embrace or kiss their pet turtles may run the
risk of catching the disease, and such close contact should there
fore be discouraged. Certainly the “dry bowl” method of raising
baby turtles, whether or not required by law, has nothing to recom
mend it and should not be used. The present legislation restricting
sales of baby turtles, while doubtless carrying a humanitarian side
benefit in saving thousands of turtles from a premature death in
the hands of youthful owners, is not justified on health grounds
since not only the substances listed above, but even domestic dogs,
cats, and birds can be sources of infection. In other words, we run
a constant risk of contracting salmonellosis whatever we do, and
that risk is only negligibly increased by keeping pet turtles.
Sea turtles in captivity are subject to various diseases, which
have been subject to detailed investigation at the Cayman Island
Turtle Farm (formerly Mariculture Ltd.). One of the most serious
of these diseases has been called LET disease because it affects
the lungs, eyes, and throat. It is first seen at an age of about four
months and is manifested by hard pulmonary lesions which may
cause the collapse of one lung with consequent asymmetrical float
ing. Vast eye lesions also develop, and the trachea becomes so
plugged with desquamated epithelium that the turtle has great dif-
847
www.Ebook777.com
ficulty breathing. This disease is caused by Clostridium botulin-
um.
Another disease of captive sea turtles has been called ‘floppy
flipper’ by the Mariculture researchers. It first appears at about
nine months and is manifested by a paralysis which starts with the
eyelids and moves down to the flippers. It is extremely contagious,
and turtles infected with it usually die by drowning.
Young sea turtles may start showing puffy skin lesions from
about two months of age onward. These lesions are caused by a
herpes-type virus and may be scratched off if no secondary bac
terial infection is present. When further infection is present, oxi
dizing agents such as potassium permanganate, silver nitrate, and
Gentian violet may be used.
Finally, Mariculture classifies some of its turtles by the acro
nym DON (dead but otherwise normal)—almost as revolting a
term as the herpetologist’s familiar DOR (dead on road).
BACKGROUND READING
“Amphibius.” Undated. Land and Water Tortoises. Water Life
Series, Dorset House, London. 32 pp.
Dowling, H. and S. Spencook. 1960. The care of pet turtles. N.Y.
Zool. Soc., 16 pp.
Frye, F.L. 1973. Husbandry, medicine and surgery in captive rep
tiles. V.M. Publishing, Inc.; Bonner Springs, Kansas. 140 pp.
Haas, R. 1967. Enjoy your turtle. Pet Library Ltd., N.Y. 32 pp.
Haga, J.B. 1972. Baby green turtles after certification! Pet/Sup-
plies/Marketing, Sept. 1972: 47-50.
Hoke, J. 1970. Turtles and their care. Franklin Watts, Inc., N.Y.
89 pp.
International Turtle and Tortoise Society Journal. 1966-1973
(numerous articles throughout all years of this journal).
Mehrtens, J. Undated. Know your turtles. Pet Library, Ltd., N.Y.
64 pp.
__________ 1974. Brief notes on the Geoemyda spinosa. Tortuga
Gazette, 10(1): 9-10.
Murphy, J.B. 1973. A review of diseases and treatment of captive
chelonians. Part 1. HISS News Journal 1(1): 5-8.
__________ 1973. A review of diseases and treatment of captive
chelonians—bacterial and viral infections. Part 2. HISS News
Journal 1(3): 77-81.
__________. 1973. A review of diseases and treatment of captive
chelonians. Protozoal infections—vectors. HISS News Journal
1(4): 123-128.
848
__________ 1973. A review of diseases and treatment of captive

chelonians: turbellarians—trematodes—cestodes—acantho-
cephalan worms—nematodes—insects—myiasis. Part 4. HISS
News Journal 1(5): 139-150.
Noel-Hume, I. and A. Noel-Hume. 1954. Tortoises, terrapins, and
turtles. W. & G. Foyle, Ltd., London. 112 pp.
Roberts, M.F. 1955. Turtles as pets. A guide to the selection and
care of turtles. T.F.H. Publications, Inc. 32 pp.
Smith, H.M. 1967. Turtles in color. T.F.H. Publications, Inc. 32 pp.
Tinkle, D.W. 1964. Playful turtles. T.F.H. Publications, Inc. 32 pp.
United States Dept, of the Interior. 1973. Convention on Inter
national Trade in Endangered Species of Wild Fauna and
Flora, signed March 3, 1973. 54 pp.
____________________. 1973. U.S. List of Endangered Foreign Fish
and Wildlife (includes all revisions through June 4, 1973).
849
www.Ebook777.com
APPENDIX I
CHECKLIST
The list below includes the names of all living and recently ex
tinct turtle species and subspecies. All subcategories are listed al
phabetically, with the exception of nominate subspecies. Syno
nyms are not given; in a few cases these may be found in the text,
but for a nearly complete listing the reader should consult Wer-
muth and Mertens 1961 (Schildkroten, Krokodile, Bruckenechsen;
Gustav Fischer, Jena). Needless to say, specialist opinion differs
on the validity of some forms and on whether certain others are
better considered species or subspecies. I have endeavored not to
be too controversial, and where specialists differ I have attempted
to make a stand on the basis of the nature of the turtles themselves
rather than on a head count of specialists in each opposing faction.
Family: Carettochelyidae (New Guinea plateless river turtles)
Carettochelys insculpta Ramsay 1887
Family: Cheloniidae (Sea turtles)

Caretta caretta (Linnaeus 1758)
Chelonia depressa Garman 1880
Chelonia mydas mydas (Linnaeus 1758)
Chelonia mydas agassizi Bocourt 1868
Eretmochelys imbricata imbricata (Linnaeus 1766)
Eretmochelys imbricata squamata Agassiz 1857
Lepidochelys kempi (Garman 1880)
Lepidochelys olivacea (Eschscholtz 1829)
Family: Chelydridae (Snapping turtles)

Chelydra serpentina serpentina (Linnaeus 1758)
Chelydra serpentina acutirostris Peters 1862
Chelydra serpentina osceola Stejneger 1918
Chelydra serpentina rossignoni (Bocourt 1868)
Macroclemys temmincki (Troost 1835)
Family: Dermatemyidae (Central American river turtles)

Dermatemys mawi Gray 1847
Family: Dermochelyidae (Leatherback turtles)

Dermochelys coriacea (Linnaeus 1766)
850
Family: Emydidae (Common freshwater turtles)

Subfamily: Emydinae
Chrysemys picta picta (Schneider 1773)
Chrysemys picta belli (Gray 1831)
Chrysemys picta dorsalis Agassiz 1857
Chrysemys picta marginata Agassiz 1857
Clemmys guttata (Schneider 1792)
Clemmys insculpta (LeConte 1830)
Clemmys marmorata marmorata (Baird and Girard 1852)
Clemmys marmorata pallida Seeliger 1945
Clemmys muhlenbergi (Schoepff 1801)
Deirochelys reticularia reticularia (Latreille 1801)
Deirochelys reticularia chrysea Schwartz 1956
Deirochelys reticularia miaria Schwartz 1956
Emydoidea blandingi (Holbrook 1838)
Emys orbicularis (Linnaeus 1758)
Graptemys barbouri Carr and Marchand 1942
Graptemys caglei Lantz 1972
Graptemys geographica (LeSueur 1817)
Graptemys flavimaculata Cagle 1954
Graptemys kohni (Baur 1890)
Graptemys nigrinoda nigrinoda Cagle 1954
Graptemys nigrinoda delticola Folkerts and Mount 1969
Graptemys oculifera (Baur 1890)
Graptemys pseudogeographica pseudogeographica (Gray 1831)
Graptemys pseudogeographica ouachitensis Cagle 1953
Graptemys pseudogeographica sabinensis Cagle 1953
Graptemys pulchra Baur 1893
Graptemys versa Stejneger 1925
Malaclemys terrapin terrapin (Schoepff 1793)
Malaclemys terrapin centrata (Latreille 1802)
Malaclemys terrapin littoralis Hay 1904
Malaclemys terrapin macrospilota Hay 1904
Malaclemys terrapin pileata (Wied 1865)
Malaclemys terrapin rhizophorarum Fowler 1906
Malaclemys terrapin tequesta Schwartz 1955
Pseudemys alabamensis Baur 1893
Pseudemys concinna concinna (LeConte 1830)
Pseudemys concinna hieroglyphica (Holbrook 1836)
Pseudemys concinna mobilensis (Holbrook 1838)
Pseudemys concinna suwanniensis Carr 1937
Pseudemys concinna texana Baur 1893
Pseudemys decorata Barbour and Carr 1940
Pseudemys decussata decussata (Gray 1831)
Pseudemys decussata angusta Barbour and Carr 1940
Pseudemys decussata plana Barbour and Carr 1940
851
www.Ebook777.com
Pseudemys felis Barbour 1935
Pseudemys floridana floridana (LeConte 1830)
Pseudemys floridana hoyi (Agassiz 1857)
Pseudemys floridana peninsularis Carr 1938
Pseudemys granti Barbour and Carr 1941
Pseudemys malonei Barbour and Carr 1938
Pseudemys nelsoni Carr 1938
Pseudemys rubriventris (LeConte 1830)
Pseudemys scripta scripta (Schoepff 1792)
Pseudemys scripta callirostris (Gray 1855)
Pseudemys scripta cataspila (Gunther 1885)
Pseudemys scripta elegans (Wied 1839)
Pseudemys scripta gaigeae Hartweg 1939
Pseudemys scripta grayi (Bocourt 1868)
Pseudemys scripta hiltoni Carr 1942
Pseudemys scripta nebulosa (Van Denburgh 1895)
Pseudemys scripta dorbignyi (Dumeril and Bibron 1835)
Pseudemys scripta ornata (Gray 1831)
Pseudemys scripta troosti (Holbrook 1836)
Pseudemys scripta venusta (Gray 1855)
Pseudemys scripta yaquia Legler and Webb 1970
Pseudemys stejnegeri Schmidt 1928
Pseudemys terrapen (Lacepede 1788)
Terrapene Carolina Carolina (Linnaeus 1758)
Terrapene Carolina bauri Taylor 1895
Terrapene Carolina major (Agassiz 1857)
Terrapene Carolina mexicana (Gray 1849)
Terrapene Carolina triunguis (Agassiz 1857)
Terrapene Carolina yucatana (Boulenger 1895)
Terrapene coahuila Schmidt and Owens 1944
Terrapene nelsoni nelsoni Stejneger 1925
Terrapene nelsoni klauberi Bogert 1943
Terrapene ornata ornata (Agassiz 1857)
Terrapene ornata luteola Smith and Ramsay 1952
Subfamily: Batagurinae
Annamemys annamensis (Siebenrock 1903)
Batagur baska (Gray 1831)
Callagur bomeoensis (Schlegel and Muller 1844)
Chinemys kwangtungensis (Pope 1934)
Chinemys megalocephala Fang 1934
Chinemys reevesi (Gray 1831)
Cuora amboinensis (Daudin 1802)
Cuora flavomarginata (Gray 1863)
Cuora galbinifrons Bourret 1939
Cuora trifasciata (Bell 1825)
Cuora yunnanensis (Boulenger 1906)
852
Cyclemys dentata (Gray 1831)

Cyclemys mouhoti Gray 1862
Geoclemys hamiltoni (Gray 1831)
Geoemyda spengleri spengleri (Gmelin 1789)
Geoemyda spengleri japonica Fan 1931
Hardella thurji (Gray 1831)
Heosemys depressa (Anderson 1875)
Heosemys grandis (Gray 1860)
Heosemys leytensis Taylor 1920
Heosemys silvatica (Henderson 1912)
Heosemys spinosa (Gray 1831)
Hieremys annandalei (Boulenger 1903)
Kachuga dhongoka (Gray 1834)
Kachuga kachuga (Gray 1831)
Kachuga smithi (Gray 1863)
Kachuga sylhetensis (Jerdon 1870)
Kachuga tecta (Gray 1831)
Kachuga tentoria tentoria (Gray 1834)
Kachuga tentoria circumdata Mertens 1969
Kachuga trivittata (Dumeril and Bibron 1835)
Malayemys subtrijuga (Schlegel and Muller 1844)
Mauremys caspica caspica (Gmelin 1774)
Mauremys caspica rivulata (Valenciennes 1833)
Mauremys japonica (Temminck and Schlegel 1833)
Mauremys leprosa (Schweigger 1812)
Mauremys mutica (Cantor 1842)
Melanochelys tricarinata (Blyth 1856)
Melanochelys trijuga trijuga (Schweigger 1812)
Melanochelys trijuga coronata (Anderson 1878)
Melanochelys trijuga edeniana Theobold 1876
Melanochelys trijuga indopeninsularis (Annandale 1913)
Melanochelys trijuga parkeri Deraniyagala 1939
Melanochelys trijuga thermalis (Lesson 1830)
Morenia ocellata (Dumeril and Bibron 1835)
Morenia petersi (Anderson 1879)
Notochelys platynota (Gray 1834)
Ocadia sinensis (Gray 1834)
Orlitia bomeensis Gray 1873
*Rhinoclemmys annulata Gray 1860
Rhinoclemmys areolata (Dumeril, Bibron and Dumeril 1851)
Rhinoclemmys diademata (Mertens 1954)
Rhinoclemmys funerea (Cope, 1876)
Rhinoclemmys melanostema (Gray 1861)
Rhinoclemmys nasuta (Boulenger 1902)
Rhinoclemmys rubida rubida (Cope 1870)
Rhinoclemmys rubida perixantha (Mosimann and Rabb 1953)
'See C.H. Ernst, 1978, Herpetologica, 31,(2) for a short review of

the genus as Callopsis.
www.Ebook777.com
Rhinoclemmys pulcherrima pulcherrima (Gray 1855)
Rhinoclemmys pulcherrima incisa (Bocourt 1868)
Rhinoclemmys pulcherrima manni (Dunn 1930)
Rhinoclemmys pulcherrima rogerbarbouri (Ernst 1978)
Rhinoclemmys punctularia (Daudin 1802)
Sacalia bealei (Gray 1831)
Siebenrockiella crassicollis (Gray 1831)
Family: Kinosternidae (American mud and musk turtles)

Claudius angustatus Cope 1865
# Kinosternon abaxillare Baur 1925
Kinosternon acutum Gray 1831

Kinosternon angustipons Legler 1965
Kinosternon bauri bauri (Garman 1891)
Kinosternon bauri palmarum Stejneger 1925
Kinosternon creaseri Hartweg 1934
Kinosternon dunni Schmidt 1947
Kinosternon flavescens flavescens (Agassiz 1857)
Kinosternon flavescens spooneri Smith 1951
Kinosternon flavescens stejnegeri Hartweg 1938
Kinosternon herrerai Stejneger 1925
Kinosternon hirtipes hirtipes (Wagler 1830)
Kinosternon hirtipes murrayi Glass and Hartweg 1951
Kinosternon integrum (LeConte 1854)
Kinosternon leucostomum
(Dumeril, Bibron, and Dumeril 1851)
Kinosternon scorpioides scorpioides (Linnaeus 1766)
Kinosternon scorpioides albogulare
(Dumeril and Bocourt 1870)
Kinosternon scorpioides carajasensis da Cunha 1970
Kinosternon scorpioides cruentatum
(Dumeril, Bibron and Dumeril 1851)
Kinosternon scorpioides pachyurum
Muller and Hellmich 1936
Kinosternon scorpioides seriei Freiberg 1936
Kinosternon sonoriense (LeConte 1854)
Kinosternon subrubrum subrubrum (Lacepede 1788)
Kinosternon subrubrum hippocrepis Gray 1856
Kinosternon subrubrum steindachneri (Siebenrock 1906)
Staurotypus salvini Gray 1864
Staurotypus triporcatus (Wiegmann 1828)
Stemotherus carinatus (Gray 1856)
Stemotherus minor minor (Agassiz 1857)
Stemotherus minor depressus Tinkle and Webb 1955
Stemotherus minor peltifer Smith and Glass 1947
Stemotherus odoratus (Latreille 1801)
854 *See main text for changes and additions.

Family: Platysternidae (Big-headed turtles)

Platystemon megacephalum megacephalum Gray 1831
Platystemon megacephalum peguense Gray 1870
Platystemon megacephalum vogeli Wermuth 1969
Family: Testudinidae (Tortoises)

Acinixys planicauda (Grandidier 1867)
Chersine angulata (Schweigger 1812)
Geochelone (Geochelone) elegans (Schoepff 1795)
Geochelone (Geochelone) pardalis pardalis (Bell 1828)
Geochelone (Geochelone) pardalis babcocki
(Loveridge 1935)
Geochelone (Geochelone) platynota (Fiyth 1863)
Geochelone (Geochelone) sulcata (Mil'er 1779)
Geochelone (Aldabrachelys) gigantea Schweigger 1812)
t Geochelone (Aldabrachelys) gouffei (F.othschild 1906)
t Geochelone (Aldabrachelys) grandidieri (Vaillant 1885)
t Geochelone (Aldabrachelys) sumeirei (Sauzier 1892)
t Geochelone (Asterochelys) abrupta (Milne-Edwards 1868)
Geochelone (Asterochelys) radiata (Shaw 1802)
Geochelone (Asterochelys) yniphora (Vaillant 1885)
Geochelone (Chelonoidis) carbonaria (Spix 1824)
Geochelone (Chelonoidis) chilensis (Gray 1870)
Geochelone (Chelonoidis) denticulata (Linnaeus 1766)
Geochelone (Chelonoidis) elephantopus elephantopus
(Harlan 1827)
Geochelone (Chelonoidis) elephantopus abingdoni
(Gunther 1877)
Geochelone (Chelonoidis) elephantopus becki
(Rothschild 1901)
Geochelone (Chelonoidis) elephantopus chathamensis
(Van Denburgh 1907)
Geochelone (Chelonoidis) elephantopus darwini
(Van Denburgh 1907)
Geochelone (Chelonoidis) elephantopus ephippium
(Gunther 1875)
t Geochelone (Chelonoidis) elephantopus galapagoensis
(Baur 1889)
Geochelone (Chelonoidis) elephantopus guntheri
(Baur 1889)
Geochelone (Chelonoidis) elephantopus hoodensis
(Van Denburgh 1907)
Geochelone (Chelonoidis) elephantopus microphyes
(Gunther 1875)
Geochelone (Chelonoidis) elephantopus phantastica
(Van Denburgh 1907)
(A f indicates a recently extinct form). 855
www.Ebook777.com
Geochelone (Chelonoidis) elephantopus porteri
(Rothschild 1903)
Geochelone (Chelonoidis) elephantopus vandenburghi
(de Sola 1930)
t Geochelone (Megalochelys) gadowi (Van Denburgh 1914)
t Geochelone (Cylindraspis) indica (Perrault 1783)
t Geochelone (Cylindraspis) inepta (Gunther 1875)
t Geochelone (Cylindraspis) vosmaeri (Schoepff 1792)
Geochelone (Indotestudo) elongata (Blyth 1853)
Geochelone (Indotestudo) travancorica (Boulenger 1907)
Geochelone (Manouria) emys (Schlegel and Muller 1844)
Geochelone (Manouria) impressa (Gunther 1882)
Gopherus agassizi (Cooper 1863)
Gopherus berlandieri (Agassiz 1857)
Gopherus flavomarginatus Legler 1959
Gopherus polyphemus (Daudin 1802)
Homopus areolatus (Thunberg 1787)
Homopus boulengeri Duerden 1906
Homopus femoralis Boulenger 1888
Homopus signatus (Walbaum 1782)
Kinixys belliana belliana Gray 1831
Kinixys belliana nogueyi (Lataste 1886)
Kinixys erosa (Schweigger 1812)
Kinixys homeana Bell 1827
Malacochersus tomieri (Siebenrock 1903)
Psammobates geometricus (Linnaeus 1758)
Psammobates oculifer (Kuhl 1820)
Psammobates tentorius tentorius (Bell 1828)
Psammobates tentorius trimeni (Boulenger 1886)
Psammobates tentorius verroxii (Smith 1839)
Pyxis arachnoides Bell 1827
Testudo (Testudo) graeca graeca Linnaeus 1758
Testudo (Testudo) graeca ibera Pallas 1814
Testudo (Testudo) graeca terrestris Forskal 1775
Testudo (Testudo) graeca zarudnyi Nikolskij
Testudo (Testudo) hermanni hermanni Gmelin 1789
Testudo (Testudo) hermanni robertmertensi Wermuth 1952
Testudo (Testudo) horsfieldi Gray 1844
Testudo (Testudo) marginata Schoepff 1792
Testudo (Pseudotestudo) kleinmanni Lortet 1883
Family: Trionychidae (Soft-shelled turtles)

Subfamily: Cyclanorbinae
Cyclanorbis elegans (Gray 1869)
Cyclanorbis senegalensis (Dumeril and Bibron 1835)
Cycloderma aubryi (Dumeril 1856)
856
Cycloderma frenatum Peters 1854

Lissemys punctata punctata (Lacepede 1788)
Lissemys punctata granosa (Schoepff 1801)
Lissemys punctata scutata (Peters 1868)
Subfamily: Trionychinae
Chitra indica (Gray 1831)
Pelochelys bibroni (Owen 1853)
Trionyx ater Webb and Legler 1960
Trionyx cartilagineus (Boddaert 1770)
Trionyx euphraticus (Daudin 1802)
Trionyx ferox (Schneider 1783)
Trionyx formosus Gray 1869
Trionyx gangeticus Cuvier 1825
Trionyx hurum Gray 1831
Trionyx leithi Gray 1872
Trionyx muticus muticus LeSueur 1827
Trionyx muticus calvatus Webb 1959
Trionyx nigricans Anderson 1875
Trionyx sinensis sinensis Wiegmann 1835
Trionyx sinensis tuberculatus Cantor 1842
Trionyx spinifer spinifer LeSueur 1827
Trionyx spinifer asper (Agassiz 1857)
Trionyx spinifer emoryi (Agassiz 1857)
Trionyx spinifer guadalupensis Webb 1962
Trionyx spinifer hartwegi (Conant and Goin 1948)
Trionyx spinifer pallidus Webb 1962
Trionyx steindachneri Siebenrock 1906
Trionyx swinhonis Boulenger 1889
Trionyx subplanus Geoffroy 1809
Trionyx triunguis (Forskal 1775)
Suborder: Pleurodira (Side-necked turtles)

Family: Chelidae
Chelodina expansa Gray 1857
Chelodina longicollis (Shaw 1802)
Chelodina novaeguineae Boulenger 1888
Chelodina oblonga Gray 1841
Chelodina parkeri Rhodin and Mittermeier 1976
Chelodina rugosa Ogilby 1890
Chelodina siebenrocki Werner 1901
Chelodina steindachneri (Schneider 1783)
Chelus fimbriatus (Schneider 1783)
Elseya dentata (Gray 1863)
Elseya latistemum Gray 1867
857
www.Ebook777.com
Elseya novaeguineae (Meyer 1874)
Emydura australis australis (Gray 1841)
Emydura australis albertisii Boulenger 1888
Emydura australis subglobosa (Krefft 1876)
Emydura kreffti (Gray 1871)
Emydura macquarrii macquarrii (Gray 1831)
Emydura macquarrii signata Ahl 1932
Hydromedusa maximiliani (Mikan 1820)
Hydromedusa tectifera Cope 1869
Phrynops (Phrynops) geoffroanus geoffroanus
(Schweigger 1812)
Phrynops (Phrynops) geoffroanus tuberosus (Peters 1870)
Phrynops (Phrynops) hilarii (Dumeril and Bibron 1835)
Phrynops (Phrynops) hogei Mertens 1967
Phrynops (Phrynops) rufipes (Spix 1824)
Phrynops (Batrachemys) dahli Zangerl and Medem 1958
Phrynops (Batrachemys) nasuta nasuta (Schweigger 1812)
Phrynops (Batrachemys) nasuta wermuthi Mertens 1970
Phrynops (Batrachemys) tuberculata (Luederwaldt 1926)
Phrynops (Batrachemys) vanderhaegei Bour 1973
Phrynops (Mesoclemmys) gibba (Schweigger 1812)
Platemys pallidipectoris Freiberg 1945
Platemys platycephala (Schneider 1792)
Platemys radiolata radiolata (Mikan 1820)
Platemys radiolata spixi Dumeril and Bibron 1835
Pseudemydura umbrina Siebenrock 1901
Family: Pelomedusidae
Erymnochelys madagascariensis (Grandidier 1867)
Pelomedusa subrufa subrufa (Lacepede 1788)
Pelomedusa subrufa olivacea (Schweigger 1812)
Peltocephalus tracaxa Dumeril and Bibron 1835
Pelusios adansoni (Schweigger 1812)
Pelusios bechuanicus Fitzsimons 1932
Pelusios carinatus Laurent 1956
Pelusios castaneus castaneus (Schweigger 1812)
Pelusios castaneus chapini Laurent 1965
Pelusios castaneus derbianus (Gray 1844)
Pelusios castaneus rhodesianus Hewitt 1927
Pelusios gabonensis (Dumeril 1856)
Pelusios nanus Laurent 1956
Pelusios niger (Dumeril and Bibron 1835)
Pelusios simmtus (Smith 1838)
Pelusios subniger (Lacepede 1788)
Pelusios williamsi williamsi Laurent 1965
Pelusios williamsi lutescens Laurent 1965
858
Podocnemis erythrocephala (Spix 1824)

Podocnemis expansa (Schweigger 1812)
Podocnemis lewyana Dumeril 1852
Podocnemis sextuberculata Comalia 1849
Podocnemis unifilis Troschel 1848
Podocnemis vogli Muller 1935
It should be noted that in some cases the usage of a name in

this listing reflects common rather than strictly correct usage. For
example, in this book it is argued that the Celebes tortoise {Geo-
chelone forsteni) is based on introduced specimens of the Travan-
core tortoise (Geochelone travancorica). Strictly speaking, there
fore, the older name (forsteni) should be used for the Travancore
tortoise. Also, rigid application of the rules of nomenclature would
oblige us to declare Geochelone elephantopus abingdoni a syno
nym of G. elephantopus ephippium, and a new name would then
have to be proposed for the Duncan tortoise (usually called ephip
pium). The Alcedo tortoise too should be re-named, the name van-
denburghi being unsatisfactory for several reasons.
Sacalia bealei should really be called Cistuda bealei, since Cis-
tuda is not a homonym of Cistudo.
Several species (notably Chelonia mydas, Eretmochelys im-
bricata, and Pseudemys scripta) certainly include more subspe
cies than are given in the list, but these additional subspecies have
yet to be named.
859
www.Ebook777.com
APPENDIX II
TURTLE GENERA
Some years ago, Kurt Vonnegut wrote a review of the Random
House Dictionary in which he pointed out that anyone with suffi
cient determination could write the best dictionary ever simply by
being certain to have all the words that the other dictionaries have,
then adding words which have joined the language since the others
were published, and then avoiding mistakes which the earlier dic
tionaries caught particular hell for. This is in essence the proce
dure I have followed in compiling the list below. While I believe the
compendium of turtle generic names given here is the most com
plete available, I freely acknowledge the pioneering work of the
late Prof. A.S. Romer in his systematic listing of reptilian genera.
As a starting point for this list, I consulted the classification sec
tion in Romer’s Osteology of the Reptiles, arranged the names in
alphabetical order, made minor changes in accordance with the
system of classification used in this book, made good the few
names omitted by Romer that had come to my attention, and
added those generic names that have been proposed since Romer’s
book was published in 1956.
This list includes names of both living and extinct genera; the
latter are preceded by a t- Valid names are given in capitals; in
valid names in lower case. Subgeneric names are given in paren
theses. In all cases, author’s name and date of publication follow
the name of the genus. In the case of invalid genera, this is fol
lowed by the valid or preferred name. For valid genera, the fami
ly, geologic periods from which the genus has been recorded, and
gross distribution are given. Abbreviations are as follows:
1) Trias.: Triassic. 2) Jur.: Jurassic. 3) Cret.: Cretaceous. 4)
Paleoc.: Paleocene. 5) Eoc.: Eocene. 6) Olig.: Oligocene. 7)
Mioc.: Miocene. 8) Plioc.: Pliocene. 9) Pleist.: Pleistocene. 10)
R.: Recent. 11) Af.: Africa. 12) As.: Asia. 13) Aust.: Australia.
14) CA.: Central America. 15) EAs.: Eastern Asia. 16) E.Ind.:
East Indies (Indonesia). 17) Eu.: Europe. 18) NA.: North Ameri
ca. 19) NAf.: North Africa. 20) Oc.: Ocean(s). 21) SA.: South
America. 22) SAs.: southern Asia. 23) W.Ind.: West Indies.
Acanthochelys Gray 1873. Syn. Platemys.

t Achelonia Wagner 1860. Syn. Eurystemum.
860
t ACHERONTEMYS Hay 1899. Chelydridae. Mioc. NA.

t ACHILEMYS Hay 1908. ?Testudinidae. Eoc. NA.
t Acichelys Meyer 1854. Syn. Eurystemum.
ACINIXYS Siebenrock 1902. Testudinidae. R. Madagascar.
Acrohydraspis Pusconi 1934. Syn. Phrynops.
Actinemys Agassiz 1857. Syn. Clemmys.
Adelochelys Baur 1896. Syn. Orlitia.
t ADOCUS Cope 1868. Dermatemyidae. U.Cret. Paleoc. NA., Eoc.
EAs.
t AGOMPHUS Cope 1871. Dermatemyidae. U.Cret. NA.
Agrione Khozatsky & Mlynarski 1966. Syn. Testudo.
t AGRIONEMYS Ckhikvadze 1970. Testudinidae. Plioc. As.
t AKROCHELYS Bergounioux 1935. ?Carettochelyidae. Eoc. Eu.
t ALAMOSEMYS Hay 1908. ?Dermatemyidae. Eoc. NA.
(ALDABRACHELYS) Loveridge & Williams 1957. Testudinidae,
Geochelone. R. Ind. Oc.
t ALLAEOCHELYS Noulet 1867. Carettochelyidae. Eoc. Eu.
t ALLOPLEURON Baur 1888. Cheloniidae. U.Cret. Eu.
t Amblypeza Hay 1908. Syn. Taphrosphys.
t Amphiemys Cope 1878. Syn. Agomphus.
Amyda Oken 1816. Syn. Trionyx.
t ANAPHOTIDEMYS Hay 1905. ?Thalassemyidae. U.Jur. Eu.
ANNAMEMYS Bourret 1939. Emydidae. R. SAs.
t ANOSTEIRA Leidy 1871. Carettochelyidae. Paleoc.-Olig. Eu.,
Eoc. NA., As.
t Anostira Lydekker 1889. Syn. Anosteira.
Anota Gray 1863. ?Syn. Pelusios.
t ANTHRACOCHELYS Bergounioux 1953. Pelomedusidae. Eoc.
Eu.
Apalone Rafinesque 1832. Syn. Trionyx.
t APEROTEMPORALIS Stromer 1934. Aperotemporalidae.
U.Cret. NAf.
t Apholidemys Pomel 1847. ?Syn. Anosteira.
t Aplax Meyer 1843. Syn. Eurystemum.
t APODICHELYS Price 1954. Pelomedusidae. Cret. SA.
t ARCHAEOCHELYS Lydekker 1889. Pleurostemidae. L.Cret.
Eu.
t Archaeochelys Bergounioux 1938. Non-testudinate. Perm. Eu.
t ARCHELON Wieland 1896. Protostegidae. U.Cret. NA.
t ARGILLOCHELYS Lydekker 1889. Cheloniidae. Eoc. Eu.
Aromochelys Gray 1855. Syn. Stemotherus.
Aspideretes Hay 1904. Syn. Trionyx.
Aspidochelys Gray 1860. Syn. Cycloderma.
Aspidonectes Wagler 1830. Syn. Trionyx.
Aspilus Gray 1864. Syn. Trionyx.
861
www.Ebook777.com
(ASTEROCHELYS) Gray 1873. Testudinidae, Geochelone.
Pleist.-R. Madagascar.
t ATLANTOCHELYS Agassiz 1849. ?Desmatochelyidae. U.Cret.
NA.
Aulacochelys Lydekker 1889. Syn. Trionyx.
Axestemys Hay 1899. Syn. Trionyx.
Axestus Cope 1872. Syn. Trionyx.
t BAENA Leidy 1870. Baenidae. U.cret.-Eoc. NA.
t BAICALEMYS Khosatzky 1970. Incertae sedis. Mioc. As.
Baikiea Gray 1870. Syn. Cyclanorbis.
t Bantuchelys Dollo 1925. Syn. Taphrosphys.
t BAPTEMYS Leidy 1870. Dermatemyidae. Eoc. NA.
Bartlettia Gray 1871. Syn. Podocnemis.
t BASILEMYS Hay 1902. Dermatemyidae. U.Cret.-Eoc. NA.
BATAGUR Gray 1855. Emydidae. R. SAs.
Batagurella Gray 1870. Syn. Kachuga.
(BATRACHEMYS) Stejneger 1909. Chelidae, Phrynops. R. SA.
Bellemys Williams 1950. Syn. Pyxis.
Bellia Gray 1869. Syn. Siebenrockiella.
f BOREMYS Lambe 1906. Neurankylidae. U.Cret. NA.
t BOTHREMYS Leidy 1865. Pelomedusidae. U.Cret. NA.
t Brodiechelys Nopsca 1928. Syn. Plesiochelys.
t BROILIA Bergounioux 1932. ?Emydidae. Olig. Eu.
Brookia Bartlett 1896. Syn. Orlitia.
t BRYOCHELYS van Beneden 1871. ?Cheloniidae. Olig. Eu.
Bysmachelys Johnston 1937. Syn. Gopherus.
t Bystra Hay 1916. Syn. Floridemys.
Cachuga Lydekker 1889. Syn. Kachuga.
t CALCARICHELYS Zangerl 1953. Protostegidae. U.Cret. NA.
Calemys Agassiz 1857. Syn. Clemmys.
CALLAGUR Gray 1870. Emydidae. R. SAs.
Callichelys Gray 1863. Syn. Pseudemys.
Callinia Gray 1869. Syn. Trionyx.
Callopsis Gray 1870. Syn. Rhinoclemmys.
t CALOCHELYS Bergounioux 1953. Pelomedusidae. Eoc. Eu.
Cantorella Gray 1870. Syn. Callagur.
Caouana Cocteau 1838. Syn. Caretta.
Caouana Gray 1844. Syn. Lepidochelys.
CARETTA Rafinesque 1814. Cheloniidae. Cret. Eoc. Eu., Eoc.
NAf., Plio. Eu., Pleist. NA., R.Oc.
Caretta Ritgen 1828. Syn. Eretmochelys.
CARETTOCHELYS Ramsay 1886. Carettochelyidae. Mioc.-R.
New Guinea.
t CAROLINOCHELYS Hay 1923. Cheloniidae. Olig. NA.
t CARTEREMYS Williams 1953. Pelomedusidae. Eoc. EAs.
Caryoderma Cope 1886. Syn. Geochelone.
862
t Castresia Stefano 1902. Syn.Anosteira.

t CATAPLEURA Cope 1870. Cheloniidae. U.Cret. NA.
Cathaiemys Lindholm 1931. Syn. Mauremys.
t (CAUDOCHELYS) Auffenberg 1963. Testudinidae, Geochelone.
Mioc.-Pleist. NA.
t CAUTLEYA Theobald 1879. Incertae sedis. Plioc. EAs.
Centrochelys Gray 1872. Syn. Geochelone.
Cephalochelys Gray 1873. Syn. Lepidochelys.
Ceramopelta Heude 1880. Syn. Trionyx.
Chaibassia Theobald 1876. Syn. Melanochelys.
t CHANGISAURUS Young 1959. Testudinidae.
t Charitemys Hay 1908. Syn. Neurankylus.
t CHEIROGASTER Bergounioux 1935. ?Testudinidae. Eoc. Eu.
Cheliphus Rafinesque 1832. Syn. Pelusios.
Cheliurus Rafinesque 1832. Syn. Chelydra.
CHELODINA Fitzinger 1826. Chelidae. Pleist.-R. Aust.
Chelomedusa Gray 1873. Syn. Hydromedusa.
Chelona Fleming 1922. Syn. Chelonia.
Chelone Brongniart 1805. Syn. Chelonia.
t Chelonemys Jourdan 1862. ?Syn. Idiochelys.
Chelonemys Gray 1864. Syn. Podocnemis.
CHELONIA Brongniart 1800. Cheloniidae. U.Cret. Eu., ?Eoc.-
Mioc. NA., R. Oc.
Chelonias Rafinesque 1814. Syn. Chelonia.
t Chelonides Maack 1869. Syn. Anaphotidemys.
(CHELONOIDIS) Fitzinger 1835. Testudinidae, Geochelone.
Mioc.-R. CA., SA.
Chelonura Fleming 1822. Syn. Chelydra.
Chelopus Rafinesque 1832. Syn. Rhinoclemys.
t CHELOSPHARGIS Zangerl 1953. Protostegidae. U.Cret. NA.
CHELUS Dumeril 1805. Chelidae. Tert.-R. SA.
t CHELYCARAPOOKUS Warren 1969. Chelycarapookidae
L.Cret., Aust.
CHELYDRA Schweigger 1812. Chelydridae. Pleist. NA., R. NA.,
SA.
Chelydra Rafinesque 1815, Syn. Chelus.
t CHELYDROPS Matthew 1924. Chelydridae. Mioc. NA.
t CHELYDROPSIS Peters 1858. ?Chelydridae. Olig.-Mioc. Eu.
Chelydura Oudemans 1894. Syn. Chelodina.
Chelymys Gray 1844. Syn. Emydura.
t CHELYOPSIS Smets 1887. Cheloniidae. Olig. Eu.
Chelymys Gray 1844. Syn. Emydura.
Chelyodina Agassiz 1846. Syn. Chelodina.
Chelyra Rafinesque 1832. Syn. Dermochelys.
Chelys Oppel 1811. Syn. Chelus.
863
www.Ebook777.com
t CHELYTHERIUM v. Meyer 1863. ?Proganochelyidae. U Trias
Eu.
t CHELYZOON v. Huene 1902. Incertae sedis. M. Trias. Eu.
Chemelys Rafinesque 1832. Syn. Trionyx.
t CHENGYUCHELYS Young and Chow 1953. Baenidae. Jur.-
Cret., China.
Chersina Gray 1831. Syn. Chersine.
CHERSINE Merrem 1820. Testudinidae. R. Af.
Chersinella Gray 1870. Syn. Psammobates.
Chersobius Fitzinger 1835. Syn. Homopus.
Chersus Wagler 1830. Syn. Testudo.
CHINEMYS Smith 1931. Emydidae. R. SAs.
t CHISTERNON Leidy 1872. Baenidae. Eoc. NA.
CHITRA Gray 1844. Trionychidae. Pleist.-R. SAs.
t CHITRACEPHALUS Dollo 1884. Aperotemporalidae. Jur or L
Cret. Eu.
Chloremys Gray 1871. Syn. Dermatemys.
CHRYSEMYS Gray 1844. Emydidae. ?Eoc.-?Mioc. Eu Plio -R
NA, R. CA, SA.
t CIMOCHELYS Owen 1842. ?Thalassemyidae. L.Cret. Eu.
t Cimoliochelys Woodward 1888. Syn. Cimochelys.
Cinctistemum Heude 1880. Syn. Trionyx.
Cinixys Wagler 1830. Syn. Kinixys.
Cinostemon Wagler 1830. Syn. Kinosternon.
Cinostemum Agassiz 1846. Syn. Kinosternon.
Cinothorax Fitzinger 1835. Syn. Kinixys.
Cistoclemmys Gray 1863. Syn. Cyclemys.
Cistuda Fleming 1822. Syn. Sacalia, Emys.
Cistudo Say 1825. Syn. Terrapene.
CLAUDIUS Cope 1865. Kinosternidae. R. CA.
t CLEMMYDOPSIS Boda 1927. Emydidae. Plio. Eu.
CLEMMYS Rotgen 1828. Emydidae. Paleoc.-R. NA.
Coelognathus Heude 1880. Syn. Trionyx.
t Colossochelys Falconer & Cautley 1844. Syn. (Megalochelys).
Colpochelys Garman 1880. Syn. Lepidochelys.
t COMPSEMYS Leidy 1856. Baenidae. U.Cret.-Palaeoc. NA.
Conchochelys Hay 1905. Syn. Trionyx.
Coptopelta Heude 1880. Syn. Trionyx.
Coriudo Fleming 1822. Syn. Dermochelys.
t CORSOCHELYS Zangerl 1960. Cheloniidae. U.Cret. NA.
t COSMOCHELYS Andrews 1919. Dermochelyidae. Eoc. Af.
t CRASPEDOCHELYS Rutimeyer 1873. Plesiochelyidae. U.Jur
Eu.
t Crassachelys Moody 1972. Syn. Euclastochelys.
t CRATOCHELONE Longman 1915. TDesmatochelyidae. L.Cret.
Aust.
864
t CROSSOCHELYS Simpson 1937. Meiolaniidae. Eoc. SA.

Cryptopus Dumeril & Bibron 1835. Syn. Lissemys.
t CTENOCHELYS Zangerl 1953. Toxochelyidae. U.Cret. Na.
Cuchoa Gray 1869. Syn. Kachuga.
CUORA Gray 1855. Emydidae. R. SAs.
t CYAMODUS Kuhn-Schnyder 1965. Incertae sedis. Mid. Trias.
Eu.
CYCLANORBIS Gray 1854. Trionychidae. R. Af.
Cyclanosteus Gray 1856. Syn. Cyclanorbis.
CYCLEMYS Bell 1834. Emydidae. R. SAs., E. Ind.
t CYCLOCHELYS Bergounioux 1953. Pelomedusidae.
CYCLODERMA Peters 1854. Trionychidae. Mioc.-R. Af.
f CYCLOPOCHELYS Goeldi 1905. Chelidae. Tert. SA.
t (CYLINDRASPIS) Fitzinger 1835. Testudinidae, Geochelone.
Subrecent, Mascarenes.
t CYMATHOLCUS Clark 1932. Testudinidae. Eoc. NA.
t CYNOCERCUS Cope 1872. ?Toxochelidae. U.Cret. NA.
t CYRTURA Jaekel 1904. Incertae sedis. ?U. Jur. Eu.
Cystoclemmys Gray 1870. Syn. Cyclemys.
f DACOCHELYS Lydekker 1889. Incertae sedis. Eoc. Eu.
t DACQUEMYS Williams 1954. Pelomedusidae.
Damonia Gray 1869. Syn. Malayemys.
DEIROCHELYS Agassiz 1857. Emydidae. Pleist.-R. NA.
DERMATEMYS Gray 1847. Dermatemyidae. R. CA.
Dermatochelis LeSueur 1829. Syn. Dermochelys.
Dermatochelys Wagler 1832. Syn. Dermochelys.
Dermochelis LeSueur 1829. Syn. Dermochelys.
DERMOCHELYS Blainville 1816. Dermochelyidae. Mioc. Eu.,
R. Oc.
t DESMATOCHELYS Williston 1894. Desmatochelyidae. U.Cret.
NA.
t DESMEMYS Wegner 1911. Desmemydinae. L.Cret. Eu.
Devisia Ogilby 1905. Syn. Chelydra.
Dhongoka Gray 1870. Syn. Kachuga.
Didicla Rafinesque 1832. Syn. Terrapene.
t Digerrhum Cope 1870. Syn. Pleurostemon.
Dirochelys Bauer 1889. Syn. Deirochelys.
t DITHYROSTERNON Pictet 1857. ?Emydidae. Olig. Eu.
Dogania Gray 1844. Syn. Trionyx.
t DOLLOCHELYS Zangerl 1971. Toxochelyidae. Eoc. Eu.
Dongoka Gray 1869. Syn. Kachuga.
(Dumerilia) Grandidier 1867. Syn. Erymnochelys .
f ECHMATEMYS Hay 1906. Emydidae. Eoc. NA.
Elephantopus Gray 1874. Syn. Geochelone.
t ELOCHELYS Nopsca 1931. Pelomedusidae. U.Cret. Eu.
ELSEYA Gray 1867. Chelidae. R. Aust.
865
www.Ebook777.com
Emia Gray 1870. Syn. Kachuga.
Emmenia Gray 1870. Syn. Clemmys.
Emyda Rafinesque 1815. Syn. Emys.
Emyda Gray 1831. Syn. Lissemys.
Emydes Brongniart 1806. Syn. Emys.
EMYDOIDEA Gray 1870. Emydidae. R. NA.
Emydoides Gray 1844. Syn. Terrapene.
Emydosaurus Agassiz 1846. Syn. Chelydra.
EMYDURA Bonaparte 1836. Chelidae. Pleist.-R. Australia.
Emyoides Gray 1844. Syn. Terrapene.
EMYS Dumeril 1806. Emydidae. ?Eoc.-R. Eu., R. NAf., As.
Emysaurus Dumeril & Bibron 1835. Syn. Chelydra.
t Enaliosaurus Seeley 1869. Syn. Thalassemys.
t EOCHELONE Dollo 1903. Cheloniidae. Eoc. Eu.
t EOSPHARGIS Lydekker 1889. ?Dermochelyidae. Eoc. Eu.
t EPIEMYS Yeh 1963. Emydidae. Plioc. As.
Eremonia Gray 1873. Syn. Caretta.
Eretmochelis Tschudi 1846. Syn. Eretmochelys.
ERETMOCHELYS Fitzinger 1843. Cheloniidae. R. Oc.
t ERGILEMYS Okhikvadze 1972. Testudinidae. Olig.-Mioc. As.
Eritmochelys Tschudi 1846. Syn. Eretmochelys.
t E RQUE LINNE SI A Dollo 1887. Toxochelyidae. Eoc. Eu.
Eryma Gray 1870. Syn. Clemmys.
ERYMNOCHELYS Baur 1888. Pelomedusidae. R. Madagascar.
Euamyda Stejneger 1944. Syn. Trionyx.
t EUBAENA Hay 1908. Eubaenidae. U.Cret. NA.
Euchelonia Tschudi 1846. Syn. Chelonia.
Euchelymys Gray 1871. Syn. Emydura.
Euchelys Girard 1858. Syn. Chelonia.
t Euclastes Cope 1867. Syn. Rhetechelys.
t EUCLASTOCHELYS Bergounioux 1955. Pelomedusidae. Eoc.
Af.
Eupachelys Leidy 1877. Syn. Geochelone.
t Euryaspis Wagner 1859. Syn. Eurystemum.
t EURYCEPHALOCHELYS Moody and Walker 1970. Trionychi-
chidae. Eoc. England.
t EURYSTERNUM Meyer 1859. Thalassemyidae. U.Jur.Eu.
t EUSARKIA Bergounioux 1952. Pelomedusidae. Eoc. Af.
t EUSTATOCHELYS. TPelomedusidae.
t FLORIDEMYS Williams 1950. Testudinidae. Plioc. NA.
Fordia Gray 1869. Syn. Trionyx.
t GAFSACHELYS de Stefano 1903. Pelomedudidae. Eoc. NAf.
t (GEISELEMYS) Khozatsky and Mlynarski 1966. Emydidae,
Geoemyda.
866
GEOCHELONE Fitzinger 1835. Testudinidae. Eoc.-Pleist. NA.,

Eoc.-R. Af., As., Mioc.-R. SA.
Geoclemmys Gray 1860. Syn. Rhinoclemys, Clemmys.
GEOCLEMYS Gray 1855. Emydidae. Eoc., R. SAs., E.Ind.
GEOEMYDA Gray 1834. Emydidae. Eoc.-R. As., R. E.Ind.
Geoemys Bonaparte 1838. Syn. Heosemys.
Geoliemys Matsumoto 1929. Syn. Mauremys.
t GLARICHELYS Zangerl 1958. Cheloniidae. Eoc.-Olig. Eu.
t GLAUCOCHELONE Dollo 1909. Cheloniidae. U.Cret. Eu.
t GLOSSOCHELYS Seeley 1871. Toxochelyidae. Eoc. Eu.
Glyptemys Agassiz 1857. Syn. Clemmys.
t GLYPTOCHELONE Dollo 1903. ?Cheloniidae. U.Cret. Eu.
t GLYPTOPS Marsh 1890. Glyptopsidae. U.Jur. Eu., U.Jur.-?U.
Cret., NA.
Gomphopelta Heude 1880. Syn. Trionyx.
Goniochelys Agassiz 1857. Syn. Stemotherus.
Goniochersus Lindholm 1929. Syn. Chersine.
Gopher Gray 1870. Syn. Gopherus.
GOPHERUS Rafinesque 1832. Testudinidae. Olig.-R. NA.
GRAPTEMYS Agassiz 1857. Emydidae. ?01ig., R. NA.
t GRAYEMYS Ckhikvadze 1970. Emydidae. Eoc. As.
Gymnopodus Dumeril and Bibron 1835. Syn. Trionyx.
Gymnopus Dumeril and Bibron 1835. Syn. Trionyx.
Gypochelys Agassiz 1857. Syn. Macroclemys.
t GYREMYS Hay 1908. ?Emydidae. U.Cret. NA.
Hadrianus Cope 1872. Syn. (Manouria).
Halichelys Fitzinger 1843. Syn. Caretta.
t HANGAIEMYS Sukhanov and Narmandakh 1974. Microbaeni-
dae Sukhanov 1964. L.Cret. As.
HARDELLA Gray 1870. Emydidae. Plioc., R. SAs.
t HAYEMYS Gaffney 1972. Baenidae. Cret. NA.
t HEISHANEMYS Bohlin 1953. Dermatemyidae. Cret. EAs.
t Helemys Rutimeyer 1859. Syn. Platychelys.
t Helochelydra Nopsca 1928. Syn. Tretostemon.
t HELOCHELYS Meyer 1854. ?Pleurostemidae. U.Cret. Eu.
Helopanoplia Hay 1908. Syn. Trionyx.
t HEMICHELYS Lydekker 1887. TCarettochelyidae. ?Eoc. SAs.
HEOSEMYS Stejneger 1902. Emydidae. R. SAs.
Heptathyra Cope 1859. Syn. Cycloderma.
Herpysmostes Gistl 1868. Syn. Eretmochelys.
t (HESPEROTESTUDO) Williams 1950. Testudinidae, Geoche-
lone. Olig.-Pleist. NA.
Heteroclemmys Peters 1874. Syn. Orlitia.
HIEREMYS Smith 1916. Emydidae. R. SAs.
HOMOPUS Dumeril & Bibron 1835. Testudinidae. R. Af.
867
www.Ebook777.com
t HOMOROPHUS Cope 1871. ?Dermatemyidae. U.Cret. NA.
t HOPLOCHELYS Hay 1908. Dermatemyidae. Paleoc.-Eoc. NA.
t HYBEMYS Leidy 1871. Emydidae. Eoc. Na.
Hydraspis Bell 1828. Syn. Chelodina.
HYDROMEDUSA Wagler 1830. Chelidae. R. SA.
Hydrone Rafinesque 1814. Syn. Emys.
t Hydropelta Meyer 1852. Syn. Eurystemum.
t HYLAEOCHELYS Lydekker 1889. Plesiochelyidae. Cret. Eu.
Ida Gray 1873. Syn. Trionyx.
t IDIOCHELYS Meyer 1839. Thalassemyidae. U.Jur. Eu.
(INDOTESTUDO) Lindholm 1929. Testudinidae, Geochelone. R.
SAs.
t ISEROSAURUS Fritsch 1905. Incertae sedis. Cret. Eu.
t ISLAMICHELYS Aslanova 1969. Cheloniidae. Olig. As.
Isola Gray 1873. Syn. Trionyx.
t ISOMETREMYS Chow & Yeh. Emydidae. Eoc. As.
Jerdonella Gray 1870. Syn. Kachuga.
KACHUGA Gray 1855. Emydidae. Plioc.-R. SAs.
t KALLOKIBOTION Nopsca 1923. Kallokibotinae. U.Cret. Eu.
t KALLISTIRA Hay 1908. ?Dermatemyidae. Eoc. NA.
t Kallokibotium Kopsca 1923. Syn. Kallokibotion.
t KANSUCHELYS Yeh 1963. Testudinidae. Eoc.-Olig. As.
KINIXYS Bell 1827. Testudinidae. R. Af.
KINOSTERNON Spix 1824. Kinosternidae. Plioc. NA., R. NA.,
CA., SA.
Kinostemum Bonaparte 1830. Syn. Kinosternon.
Kinothorax Gray 1863. Syn. Kinixys.
Kinyxis Bonaparte 1836. Syn. Kinixys.
t KUROBECHELYS Shikama 1956. Cheloniidae. Mioc. Japan.
Landemania Gray 1869. Syn. Trionyx.
t LEMBONAX Cope 1870. ?Cheloniidae. Eoc. NA.
LEPIDOCHELYS Fitzinger 1843. Cheloniidae. R. Oc.
t LEPTOCHELYS Bergounioux & Crouzel 1965. Carettochelyidae.
Sansan, Aquitaine.
Liemys Boulenger 1897. Syn. Orlitia.
Limnochelone Werner 1901. Syn. Dermatemys.
t LINDHOLMEMYS Riabinin 1935. Dermatemyidae. U.Cret. As.
LISSEMYS Smith 1931. Trionychidae. Plioc.-R. SAs.
t LOPHOCHELYS Zangerl 1953. Toxochelyidae. U.Cret. NA.
Lutremys Gray 1844. Syn. Emys.
t Lytoloma Cope 1870. ?Syn. Osteopygis.
t MACROBAENA Tatarinov 1959. Macrobaenidae. Eoc. EAs.
t MACROCEPHALOCHELYS Pidoplitshka & Tarashchuk 1960.
Platystemidae. Plioc. Ukraine.
Macrochelys Gray 1856. Syn. Macroclemys.
Macrochelys v. Beneden 1871. Syn. Psephophorus.
868
Macrochelys v. Meyer 1858. ?Syn. Geochelone.

Macroclemmys Strauch 1862. Syn. Macroclemys.
MACROCLEMYS Gray 1855. Chelydridae. Mioc.-R. NA.
Madakinixys Vuillemin 1972. Syn. Kinixys.
Malaclemmys Fowler 1906. Syn. Malaclemys.
MALACLEMYS Gray 1844. Emydidae. R. NA.
MALACOCHERSUS Lindholm 1929. Testudinidae. R. Af.
Malacoclemmys Agassiz 1857. Syn. Malaclemys.
MALAYEMYS Lindholm 1931. Emydidae. Plio.-R. SAs., E. E.
Ind.
Maloclemmys Agassiz 1846. Syn. Malaclemys.
t MANCHUROCHELYS Endo & Shikama 1942. Sinemydidae. L.
Cret. EAs.
(MANOURIA) Gray 1854. Testudinidae, Geochelone. Eoc. NA.,
R. SAs.
Matamata Merrem 1820. Syn. Chelus.
MAUREMYS Gray 1870. Emydidae. R. Eu., NAf., As.
Medaestia Wussow 1916. Syn. Testudo.
Megachersine Hewitt 1933. Syn. Geochelone.
t (MEGALOCHELYS) Falconer & Cautley 1837. Testudinidae,
Geochelone. Plios. As., Pleist. Mauritius.
Megalochelys Fitzinger 1843. Syn. Geochelone.
t Megasternon Gray 1842. ?Syn. Pleurostemon.
t Megasternum Lydekker 1889. Syn. Pleurostemon.
Megemys Gistl 1848. Syn. Chelonia.
t MEIOLANIA Owen 1886. Meiolaniidae. Pleist. Aust.
Melanemys Schuffeldt 1919. Syn. Clemmys.
MELANOCHELYS Gray 1869. Emydidae. E. SAs.
t MESOCHELYS Evans and Kemp 1975. L. Cret. Eu.
(MESOCLEMMYS) Gray 1873. Chelidae, Phrynops. R. SA.
Mesodeca Rafinesque 1832. Syn. Trionyx.
t MIOCARETTA Deraniyagala 1967. Cheloniidae. Mioc. Ceylon,
t Miolania Lydekker 1889. Syn. Meiolania.
t MIOTESTUDO Deraniyagala 1967. Testudinidae. Mioc. SAs.
t (MONACHELYS) Williams 1952. Testudinidae, Geochelone. Sub-
recent W.Ind.
t MONGOLEMYS Khosatzky and Mlynarski 1971. Cret. As.
Monoclida Rafinesque 1832. Syn. Terrapene.
MORENIA Gray 1870. Emydidae. R. SAs.
Mydas Cocteau 1838. Syn. Chelonia.
Mydasea Gervais 1843. Syn. Chelonia.
t Naiadochelys Hay 1908. ?Pelomedusidae U.Cret. NA., SA.
(nomen dubium)
Nanemys Agassiz 1857. Syn. Clemmys.
t NANHSIUNGCHELYS Young 1965. Incertae sedis. Cret. China,
t NAOMICHELYS Hay 1908. Pleurostemidae. U.Jur. NA.
869
www.Ebook777.com
Natator McCulloch 1908. Syn. Chelonia.
Nectemys Agassiz 1857. Syn. Pseudemys.
t NEOCHELYS Bergounioux 1954. Pelomedusidae. Eoc. Eu.
Neoemys Lindholm 1929. Syn. Emydoidea.
Neotestudo Hewitt 1931. Syn. Chersine.
t NEPTUNOCHELYS Wieland 1900. ?Desmatochelyidae. U.Cret.
NA.
t NEURANKYLUS Lambe 1902. Neurankylidae. U.Cret. NA.
Nicoria Gray 1855. Syn. Rhinoclemys.
Nilssonia Gray 1872. Syn. Trionyx.
t NIOLAMIA Ameghino 1899. Meiolaniidae. U.Cret. SA.
Notoa Gray 1863. Syn. Pelusios.
t NOTOCHELONE Lydekker 1889. ?Desmatochelyidae. Cret.
Aust.
NOTOCHELYS Gray 1863. Emydidae. R. SAs., E.Ind.
t Notochelys Owen 1882. Syn. Notochelone.
t NOTOEMYS Cattoi and Freiberg 1961. Plesiochelyidae. Jur. SA.
t NOTOMORPHA Cope 1872. ?Dermatemyidae. Eoc. NA.
OCADIA Gray 1870. Emydidae. Eoc. Mioc. Eu., Plioc. Egypt,
Mongolia, R. EAs.
Odontochelone de Stefano 1902. Syn. Trionyx.
Onychochelys Gray 1873. Syn. Eretmochelys.
Onychotria Gray 1849. Syn. Terrapene.
Ophichelone Jarocki 1822. Syn. Chelydra.
ORLITIA Gray 1873. Emydidae. R. SAs.
Oscaria Gray 1873. Syn. Trionyx.
t OSTEOPYGIS Cope 1868. Toxochelyidae. U.Cret. NA.
Ozotheca Agassiz 1857. Syn. Stemotherus.
t PACHYCHELYS v. Beneden 1871. Cheloniidae. L.Plioc. Eu.
t Pachyrhynchus Dollo 1886. ?Syn. Erquelinnesia.
t PALAEOCHELYS Meyer 1847. ?Emydidae. Olig. Eu., As.
t PALAEOCHELYS Valenciennes 1863. Carettochelyidae. U.Cret.
Eu.
t Palaeochelys Gray 1870. Syn. Paleaspis.
t Palaeomedusa Meyer 1860. Syn. Eurystemum.
t PALAEOTHECA Cope 1872. ?Emydidae. Eoc. NA.
Palaeotrionyx Schmidt 1945. Syn. Trionyx.
t PALATOBAENA Gaffney 1972. Baenidae. Cret.-Palaeoc. NA.
t PALEASPIS Gray 1870. Pelomedusidae. Eoc. Eu.
t Palemys Gray 1870. Syn. Paleaspis.
Pampatestudo Lindholm 1929. Syn. Geochelone.
Pangshura Gray 1855. Syn. Kachuga.
t PARACHELYS Meyer 1864. Plesiochelyidae. U.Jur. Eu.
t Parahydraspis Wieland 1923. Syn. Phrynops.
Paralichelys Bergounioux 1935. ?Syn. Mauremys.
Pariemys Cope 1865. Syn. Terrapene.
870
t PATANEMYS Andrews 1920. Dermatemyidae. Eoc. Eu.

t PEISHANEMYS Bohlin 1953. Dermatemyidae. Cret. EAs.
t PELOBATOCHELYS Seeley 1875. Thalassemyidae. U.Jur.-
L.Cret. Eu.
PELOCHELYS Gray 1864. Trionychidae. R. SAs.
t PELOCOMASTES de Vis 1897. Chelidae. Pleist. Aust.
Pelodiscus Fitzinger 1835. Syn. Trionyx.
PELOMEDUSA Wagler 1830. Pelomedusidae. Olig.-R. Af.
Peltastes Gray 1869. Syn. Psammobates.
PELTOCEPHALUS Dumeril and Bibron 1835. Pelomedusidae.
R.SA.
t Peltochelys Dollo 1884. ?Syn. Tretostemon.
Peltonia Gray 1872. Syn. Testudo.
PELUSIOS Wagler 1830. Pelomedusidae. Mioc.-R. Af.
Pelusius Agassiz 1848. Syn. Pelusios.
Pentonyx Dumeril and Bibron 1835. Syn. Pelomedusa.
t PERITRESIUS Cope 1870. Cheloniidae. U.Cret.-Mioc. NA.
PHRYNOPS Wagler 1830. Chelidae. Plioc.-R. SA.
t Phyllemys Schmidt 1944. Syn. Toxochelys.
t PLANIPLASTRON Ckhikvadze 1971. Platysternidae. Olig. As.
Plastomenus Cope 1873. Syn. Trionyx.
t Plastremys Owen 1881. Syn. Trachydermochelys.
PLATEMYS Wagler 1830. Chelidae. R. SA.
t PLATYARKIA Bergounioux 1953. Pelomedusidae.
t PLATYCHELOIDES Haughton 1928. Pelomedusidae. L.Cret.
EAf.
f PLATYCHELONE Nopsca 1928. Cheloniidae.
t PLATYCHELYS Wagner 1853. Pleurostemidae. L.-U.Jur. Eu.
t Platypeltis Fitzinger 1835. Syn. Trionyx.
PLATYSTERNON Gray 1831. Platysternidae. R. SAs.
Platystemum Agassiz 1846. Syn. Platystemon.
Platythyra Agassiz 1857. Syn. Kinosternon.
t PLESIOBAENA Gaffney 1972. Baenidae. Cret.-Palaeoc. NA.
t PLESIOCHELYS Rutimeyer 1873. Plesiochelyidae. U.Jur.-
L.Cret. Eu.
t PLEUROSTERNON Owen 1853. Pleurostemidae. U.Jur.-
L.Cret. Eu.
t Pleurostemum Lydekker 1889. Syn. Pleurostemon.
Pliochelys Portis 1891. ?Syn. Caretta.
PODOCNEMIS Wagler 1830. Pelomedusidae. U.Cret.-R. SA.,
t POLYECHMATEMYS Ping 1930. ?Emydidae. Plioc. EAs.
t Polystemon Portis 1882. ?Pelomedusidae. U.Cret.-Eoc. Eu. (no
men vanum)
t POLYTHORAX Cope 1877. ?Baenidae. U.Cret. NA.
t PORTHOCHELYS Williston 1901. Toxochelyidae. U.Cret. NA.
871
www.Ebook777.com
t PORTLANDEMYS Gaffney 1975. Plesiochelyidae. Jur. Eu.
Potamochelys Fitzinger 1843. Syn. Trionyx.
t Potamochelys Bergounioux and Crouzel 1968. ?Pelomedusidae.
Cret. At.
t PRIONOCHELYS Zangerl 1953. Toxochelyidae. U.Cret. NA.
t PROBAENA Hay 1903. ?Pleurostemidae. U.Jur. NA.
t Prochonias Cope 1870. Syn. Taphrosphys.
t PROCOLPOCHELYS Hay 1908. ?Cheloniidae. Mioc. NA.
t PROERETMOCHELYS Dollo 1903. ?Thalassemyidae. L.Cret.
Eu.
t PROGANOCHELYS Baur 1887. Proganochelyidae. U.Trias. Eu.
Proganosaurus Portis 1891. ?Syn. Caretta.
t PROMALACOCLEMMYS v. Reinach 1900. ?Emydidae. Mioc.
Eu.
t Propleura 1870. ?Syn. Osteopygis.
t PROTEMYS Owen 1851. Toxochelyidae. Eoc. Eu.
t PROTEROCHERSIS Fraas 1913. Proganochelyidae. U.Trias.
Eu.
t PROTESTUDO Ckhikvadze 1971. Testudinidae. Olig.-Plioc. As.
t PROTOCHELYS Lydekker 1889. ?Pleurosternidae. Mid. Jur.
Eu.
t PROTOSPHARGIS Capellini 1884. ?Protostegidae. U.Cret. NA.
t PROTOSTEGA Cope 1871. Protostegidae. U.Cret. NA.
PSAMMOBATES Fitzinger 1835. Testudinidae. R. At.
t Psammochelys Quenstedt 1889. Syn. Proganochelys.
t PSEPHOPHORUS v. Meyer 1847. Dermochelidae. Eoc.-Mioc.
NA., Eoc.-Plioc. Eu., Eoc. NAf.
t PSEUDANOSTEIRA Clark 1932. Carettochelyidae. Eoc. NA.,
Eu.
PSEUDEMYDURA Siebenrock 1901. Chelidae. R. Aust.
PSEUDEMYS Gray 1855. Emydidae. Olig.-R. NA., Pleist.-R.
W.Ind., R. CA., SA.
Pseudocadia Lindholm 1931. Syn. Mauremys.
Pseudomopus Hewitt 1931. Syn. Homopus.
t PSEUDOSPHARGIS Dames 1894. ?Protostegidae. Olig. Eu.
(PSEUDOTESTUDO) Loveridge and Williams 1957.
Testudinidae, Geochelone. R. Af.
t Pseudotrionyx Dollo 1886. Syn. Anosteira.
Psilognathus Heude 1880. Syn. Trionyx.
Ptychemys Agassiz 1857. Syn. Pseudemys.
t PTYCHOGASTER Pomel 1847. Ptychogastridae. Olig., Mioc.
Eu.
t PUPPIGERUS Cope 1870. Cheloniidae. Eoc. Eu.
Pyxiclemmys Gray 1863. Syn. Cuora.
+ PYGMAEOCHELYS Laube 1896. ?Thalassemyidae. U.Cret. Eu.
Pyxidea Gray 1863. Emydidae. Syn. Cyclemys.
872
Pyxidemys Fitzinger 1835. Syn. Terrapene.

PYXIS Bell 1827. Testudinidae. R. Madagascar.
Pyxoides Vuillemin & Domergue 1972. Syn. Pyxis.
Rafetus Gray 1864. Syn. Trionyx.
Rapara Gray 1825. Syn. Chelydra.
Redamia Gray 1870. Syn. Pseudemys.
t RHETECHELYS Hay 1908. Toxochelyidae. U.Cret. NA.
Rhinemys Wagler 1830. Syn. Phrynops.
Rhinemys Gunther 1865. Syn. (Mesoclemmys).
Rhinemys Boulenger 1889. Syn. Batrachemys.
t RHINOCHELYS Seeley 1869. Desmatochelyidae. L.Cret. Eu.
RHINOCLEMMYS Fitzinger 1835. Emydidae. R. CA., SA.
Rhinoclemys Gray. Syn. Rhinoclemmys.
t ROSASIA Carrington de Costa 1940. ?Pelomedusidae. U.Cret.
Eu.
t ROXOCHELYS Price 1954. Pelomedusidae. Cret. SA.
SAC ALIA Gray 1870. Emydidae. R. As.
t SAKYA Bogachev 1960. Emydidae. Plioc. As.
Sarbieria Gray 1869. Syn. Trionyx.
t SAURISCHIOCOMES Kuhn 1939. ?Proganochelyidae. U.Trias.
Eu.
Saurochelys Berthold 1827. Syn. Chelydra.
Scapia Gray 1869. Syn. Geochelone.
t SCUTEMYS Wiman 1930. ?Plesiochelyidae. L.Cret. EAs.
Scytena Gray 1834. Syn. Dermochelys.
Scytine Wagler 1830. Syn. Dermochelys.
Scytina Dumeril and Bibron 1835. Syn. Dermochelys.
t SENONEMYS Bohn 1966. Emydidae. Cret. Eu.
t SENRUYEMYS Shikama 1953. Incertae sedis. Olig.-Mioc.
Japan.
Seytina Wagler 1830. Syn. Dermochelys.
t SHANSIEMYS Yeh 1963. Emydidae. Plioc. As.
Sharemys Gilmore 1931. ?Syn. Mauremys.
t SHWEBOEMYS Swinton 1939. Pelomedusidae. Eoc. NAf., Eoc.-
Pleist. SAs.
SIEBENROCKIELLA Lindholm 1929. Emydidae. R. As.
Sinaspideretes Young and Chow 1853. Syn. Trionyx.
t SINEMYS Wiman 1930. Sinemydidae. U.Jur.-L.Cret. EAs.
t SINOCHELYS Wiman 1930. ?Dermatemyidae. L.Cret. EAs.
t SINOHADRIANUS Ping 1929. ?Testudinidae. Eoc. EAs.
Siphargis Risso 1826. Syn. Dermochelys.
t SOLNHOFIA Gaffney 1975. Family incert. U. Jur. Eu.
t SONTIOCHELYS Stache 1905. Thalassemyidae. U.Cret. Eu.
Spatulemys Gray 1872. Syn. Phrynops.
Sphargis Merrem 1820. Syn. Dermochelys.
Sphragis Agassiz 1846. Syn. Dermochelys.
873
www.Ebook777.com
(SPINEMYS) Khozatsky & Mlynarski 1966. Emydidae, Geoe-
myda.
Stauremys Gray 1864. Syn. Staurotypus.
STAUROTYPUS Wagler 1830. Kinosternidae. R. CA.
t STEGOCHELYS Jaekel 1915. Proganochelyidae. U.Trias. Eu.
t STEGOCHELYS Lydekker 1889. Thalassemyidae. U.Jur. Eu.
t STEREOGENYS Andrews 1901. Pelomedusidae. U.Eoc. NAf.,
Eoc. Eu.
Stemothaerus Bell 1825. Syn. Pelusios.
Stemothaerus Gray 1831. Syn. Pelusios.
Stemothaerus Wagler 1830. Syn. Cyclemys.
STUPENDEMYS Wood 1976. Pelomedusidae. Mioc. SA.
STERNOTHERUS Gray 1825. Kinosternidae. R. NA.
Stigmochelys Gray 1873. Syn. Geochelone.
t STYGIOCHELYS Gaffney and Hiatt 1971. Baenidae. Cret. NA.
t STYLEMYS Leidy 1851. Testudinidae. ?01ig.-Mioc. NA., ?Mioc.
Eu.
t Stylemys Maack 1869. Syn. Plesiochelys.
Swanka Gray 1844. Syn. Kinosternon.
t SYLLOMUS Cope 1896. Cheloniidae. Mioc. NA.
Tanoa Gray 1863. Syn. Pelusios.
t TAPHROSPHYS Cope 1869. Pelomedusidae. U.Cret., Mioc. NA.,
Eoc. SA., Paleoc. Af.
Teleopus LeConte 1854. Syn. Geochelone.
t TEMNOCLEMMYS Bergounioux 1961. Emydidae. Mioc. Eu.
Temnognathus Heude 1880. Syn. Trionyx.
Temnotrionyx Hay 1908. Syn. Trionyx.
TERRAPENE Merrem 1820. Emydidae. Mioc.-R. NA., R. CA.
Terrapene Bonaparte 1831. Syn. Clemmys.
Terrapenne Gistl 1848. Syn. Terrapene.
Terraphene Gray 1825. Syn. Terrapene.
Testudinella Gray 1870. Syn. Testudo.
Testudinites Weiss 1830. Syn. Geochelone.
TESTUDO Linnaeus 1758. Testudinidae. Eoc.-R. Eu., NAf., As.
Tetraonyx Gray 1831. Syn. Batagur.
Tetrathyra Gray 1865. Syn. Cyclanorbis.
t THALASSEMYS Rutimeyer 1859. Thalassemyidae. U.Jur. Eu.
Thalassiochelis Nardo 1864. Syn. Caretta.
Thalassochelys Fitzinger 1835. Syn. Caretta.
Therapene Schinz 1822. Syn. Terrapene.
t THESCELUS Hay 1908. Neurankylidae. U.Cret. NA.
t THINOCHELYS Zangerl 1953. Toxochelyidae. U.Cret. NA.
t Tholemys Andrews 1921. Syn. Plesiochelys.
Thyrosternum Agassiz 1857. Syn. Kinosternon.
TOMOCHELONE Dollo 1909. Cheloniidae. U.Cret. Eu.
Tortistemum Heude 1880. Syn. Trionyx.
Tortuga Molina 1789. Syn. Dermochelys.

Toxaspis Cope 1895. Syn. Terrapene.
t TOXOCHELYS Cope 1873. Toxochelyidae. U.Cret. NA.
Trachemys Agassiz 1857. Syn. Pseudemys.
t TRACHYASPIS Meyer 1843. ?Dermatemyidae. Eoc., Mioc. Eu.,
At.
t TRACHYDERMOCHELYS Seeley 1869. ?Pleurostemidae. L.
Cret. Eu.
t TRETOSTERNON Owen 1842. Dermatemyidae. U.Jur., L.Cret.,
Eu.
t Tretosternum Agassiz 1842. Syn. Tretosternon.
t TRIASSOCHELYS Jaekel 1918. Proganochelyidae. U.Trias. Eu.
t TRINITICHELYS Gaffney 1972. Baenidae. Cret. NA.
TRIONYX Geoffroy 1809. Trionychidae. U.Cret.-R. NA., As.,
Cret.-Plioc. Eu., Mioc.-R. Af., Pleist.-R. Indonesia,
t TROPIDEMYS Rutimeyer 1873. ?Thalassemyidae. U.Jur.-L.
Cret. Eu.
t TSAOTANEMYS Bohlin 1953. Dermatemyidae. Cret. EAs.
Tyrse Gray 1844. Syn. Trionyx.
Uronyx Rafinesque 1832. Syn. Kinosternon.
t Wincania Gray 1870. Syn. Plesiochelys.
t XENOCHELYS Hay 1906. Kinosternidae. Olig. NA.
Xerobates Agassiz 1857. Syn. Gopherus.
f YAXARTEMYS Riabinin 1948. Thalassemyidae. U.Jur. NAs.
Yuen Heude 1880. Syn. Trionyx.
t YUMENEMYS Bohlin 1953. Incertae sedis. Cret. EAs.
t ZAISANEMYS Ckhikvadze 1973. Emydidae. Olig. As.
t ZANGERLIA Mlynarski 1972. Testudinidae. U.Cret. As.
t ZYGORAMMA Cope 1870. ?Dermatemyidae. U.Cret. NA.
875
www.Ebook777.com
GLOSSARY
ABDOMINAL SCUTES — The third rearmost pair of plastral
scutes. Frequently the largest of the scutes of the plastron.
ACETABULUM — The large socket in the side of the pelvis into
which the head of the femur fits.
ACROMION — The elongate process from the scapula, which in
turtles is directed postero-mesially.
ALVEOLAR — Pertaining to the functional, biting part of the jaw.
AMPHICOELOUS — A term used to describe a vertebra with a
socket at each end.
ANAEROBIC — Not requiring oxygen.
ANAL SCUTES — The rearmost pair of plastral scutes.
ANOMALOUS — Irregular or abnormal.
ANTILLEAN - Pertaining to the West Indies.
ARCADE — A narrow bony bridge.
ARTICULAR — A small bone in the lower jaw which provides the
surface for the hinging action.
ATLAS — The first cervical vertebra, adjacent to the skull.
AXILLARY NOTCH — The notch in the front part of the shell into
which the front leg fits, and from which it protrudes.
AXILLARY SCUTES — The small scutes present in the plastron
opposite the armpit in most turtles.
AXIS — The second cervical vertebra.
BASICRANIAL — Referring to that part of the skull between the
premaxillae and the occipital condyle.
BASISPHENOID — A skull bone, often of triangular superficial
aspect, located between the basioccipital and the pterygoids,
located on the midline and toward the rear of the ventral aspect
of the skull.
BEAK — The horny covering of the jaws, usually a single plate
over each jaw surface.
BICUSPID — Having two cusps.
BIFURCATE — Divided into two branches.
BIPHYLETIC — A term used to describe a genus or higher group
of organisms descended by two independent paths from an an
cestral group.
BOSS — A knoblike swelling.
BRIDGE — The part of the shell of the turtle which connects the
carapace and the plastron.
876
CALLOSITY — A roughened area of skin, sometimes with super

ficial bone exposed.
CARAPACE — The dorsal part of the shell of a turtle.
CARUNCLE — The horny tubercle on the snout of baby turtles
used to break through the eggshell.
CAUDAL — Pertaining to the tail.
CENTRAL SCUTE — One of the (usually five) large scutes ex
tending down the midline of the carapace.
CENTRUM — The body of a vertebra, as opposed to the arches
and zygapophyses.
CERVICAL — Pertaining to the neck.
CHOANA — The opening of the nasal chambers into the roof of the
mouth.
CLAVICULAR — Pertaining to the clavicle, or collarbone.
CLINE — A gradual change in a structural or physiological char
acter occurring within a group over geographical distance or
environmental change; a series of organisms that show such a
gradual change.
CLOACA — The chamber into which the digestive, urinary, and
reproductive organs empty.
COLUMELLA — The slender bone which transmits sound from
the tympanum to the inner ear.
COLUMNAR — Column-shaped, or cylindrical.
CONDYLE — An articular protruberance, such as the occipital
condyle at the rear of the skull.
CONFLUENT — Flowing or merging together.
CORACOID — A fan-shaped bone in the shoulder girdle connected
to the scapula.
CORDIFORM — Heart-shaped.
CORNEOSCUTE — Obsolete or rare term for scute.
CORONOID — The bone forming the most dorsal part of the lower
jaw, to which the jaw-closing muscles attach.
COSTALS — The (usually four) large scutes running down each
side of the carapace.
COTYLE — A cavity into which a condyle fits.
CRETACEOUS — A geologic period extending from approximate
ly 140 million to 65 million years before the present time.
CRUCIFORM — Cross-shaped.
CRURO — Pertaining to the thigh or leg.
CRYPTOZOIC — Conducting a hidden or secluded way of life.
CUSP — A sharp projection, typically from the edge of the jaw.
DEME — An isolated population of a species.
DENTARY — The lower jawbone in which the teeth are set in
most vertebrates.
DICHOTOMOUS — Divided into two parts, as in a dichotomous
identification key, which presents two conflicting alternatives
877
www.Ebook777.com
at each stage, which when followed through lead to an unam
biguous identification.
DIGITIGRADE — A term used to describe a gait in which only the
digits are in contact with the ground.
ECOTONE — A transitional area between two distinct ecosys
tems.
EDENTULOUS — Without teeth, especially the condition of
having lost teeth once present.
ENDEMIC — Occurring naturally in and restricted to a particular
area.
ENTOPLASTRON — A single antero-centrally located plastral
bone occurring between and surrounded by the epiplastra and
the hyoplastra.
EOCENE — A geologic epoch which extended from approximate
ly 55 million to approximately 35 million years ago.
EPIPLASTRON — One of the two anteriormost bones in the turtle
plastron.
ESOPHAGUS — The tube which conveys food from the mouth to
the stomach.
ESTIVATION — State of inactivity during long periods of drought
or high temperatures.
FEMORALS — The penultimate scutes of the plastron.
FEMUR — The long bone forming the upper part of the hind leg.
FENESTRATED — Eroded through or penetrated by openings.
FIBULA — The smaller of the two parallel bones forming the
lower part of the hind leg.
FLUVIATILE — Living in rivers and streams.
FONTANELLE — An opening, especially one in the skull or shell
present only in young turtles and filled in as maturity is
reached.
FORAMEN — A permanent opening in a bone through which a
structure such as a nerve or major blood vessel passes.
FRONTAL BONE — One of two mesially contiguous bones which
make up part of the skull roof of a turtle.
GINGLYMOIDY — A condition of a vertebra in which laterally
paired convexities or concavities are present in the articular
surfaces.
GIRDLE — A bony structure located at the front (shoulder girdle)
and rear (pelvic girdle) of the body trunk which provides sup
port for the arms or forelimbs and legs or hindlimbs respective
ly-
GLENOID FOSSA — The socket in the scapula which receives the
head of the humerus.
GLOTTIS — The closable opening in the floor of the mouth into
which the windpipe opens.
GLYCOLYSIS — The biochemical process by which glycogen is
broken down to provide energy.

GULAR SCUTES — The frontmost scutes of the plastron, usually
paired but sometimes single.
HAMMOCK — An island of trees, raised above a surrounded
flooded prairie area, such as is present in the Florida Ever
glades.
HUMERAL SCUTES — The second pair of scutes in the turtle
plastron.
HUMERUS — The long bone which forms the upper part of the
forelimb.
HYOID — The bony and cartilaginous structure in the floor of the
mouth which supports the tongue.
HYOPLASTRA — The second frontmost pair of plastral bones.
HYPODIGM — The sum total of specimens available to the de-
scriber of a species.
HYPOPLASTRA — The second rearmost pair of plastral bones.
ILIAC — Pertaining to the ilium, or hip bone.
IMBRICATE — Lying overlapped, like shingles on a roof.
INFRAMARGINAL SCUTES — Scutes situated between the mar
ginals of the carapace and the pectoral and abdominal scutes of
the plastron in certain turtles.
INGUINAL NOTCH — The notch at the rear of the bridge, from
which the hind leg protrudes.
INGUINAL SCUTES — Scutes located just in front of the inguinal
notch.
INTERANAL SCUTE — A small scute occasionally present be
tween the anal scutes.
INTERFEMORAL SEAM — The seam running between the
femoral scutes.
INTERGULAR SCUTE — A small scute sometimes present which
separates, or partially separates, the gular scutes.
INTERHUMERAL SEAM — The seam running between the
humeral scutes.
INTERPECTORAL SEAM — The seam running between the pec
toral scutes.
INTERXIPHIPLASTRAL NOTCH — The median notch at the
rear of the plastron.
JUGAL — The bone which forms the postero-ventral margin of the
orbit in a turtle skull.
JURASSIC — A geologic period extending from approximately
185 million years to approximately 140 million years before the
present time.
KINESIS — Free movement, particularly between two contiguous
bones or bony structures.
LAMELLA — A thin plate, typically of bone or keratin.
LAMINA — Alternative name for scute.
879
www.Ebook777.com
LATERALS — Alternative name for costals.
MARGINAL SCUTES — The small scutes around the edge of the
carapace.
MECKEL’S CARTILAGE — A strip of cartilage located inside the
posterior parts of the lower jaw.
MELANISM — The development of excessive dark pigment.
MESENTERY — A sheet-like internal ligament with a supportive
or suspensory function.
MESIC — A term used to describe a habitat or ecosystem with an
intermediate level of moisture.
MESOPLASTRA — Plastral bones present in certain turtles; lo
cated between the hyoplastra and hypoplastra.
MESOZOIC — A geologic era, often known as the ‘Age of Rep
tiles,’ encompassing the Triassic, Jurassic, and Cretaceous
periods.
MIOCENE — A geologic epoch, extending from approximately 20
million to approximately 12 million years before the present
time.
MONOTYPIC — A taxonomic category containing only one imme
diately subordinate category, as a family with only one genus
or a genus with only one species.
NASOPALATINE FORAMEN — The opening in the upper part of
the bony palate into which the nasal chamber opens.
NEURAL BONES — The median row of bones in the turtle cara
pace.
NUCHAL BONE — The large bone at the front of the turtle cara
pace.
NUCHAL SCUTE — A small scute present in many turtles at the
front of the carapace.
OBTURATOR FORAMEN — The large central opening in the pel
vis, sometimes vertically divided.
OCCIPITAL — Pertaining to the back of the skull.
OCELLUS — A ring-shaped marking, often with a dark center.
OLIGOCENE — A geologic epoch extending from approximately
35 million years to approximately 20 million years before the
present time.
OPISTHOCOELOUS — A term used to describe a vertebra with a
posterior articular concavity.
OVIPOSITOR — A soft tubular structure extruded from the vent
during oviposition or egg-laying.
PALATINE — One of two bones making up the anterior part of the
palate of turtles; they are usually separated by the vomer.
PALEOCENE — A geologic epoch which extended from approxi
mately 65 million to approximately 55 million years before the
present time.
PAPILLA — A soft, elongate protruberance.
880
PARALLELISM — Similar evolution of unrelated organisms in

similar, but separated, environments.
PARIETAL — One of two mesially contiguous bones which make
up the posterior part of the skull roof of a turtle.
PAROCCIPITAL — A bone at the rear of the turtle skull, which
lies between the exoccipital and squamosal bones.
PECTORAL SCUTES — The third frontmost pair of plastral
scutes.
PEDICLE — A thickened area of a bone designed to withstand
weight or pressure.
PENTADACTYLY — The condition of having five digits in a hand
or foot.
PERIPHERAL BONES — The bones around the edge of the turtle
carapace.
PERMIAN — A geologic period which extended from approxi
mately 275 million to 230 million years before the present time.
PLASTRAL LOBE — That part of the plastron which lies in front
of the axillary notch or behind the inguinal notch.
PLASTRON — The ventral shell of a turtle.
PLEISTOCENE — A geologic epoch which extended from ap
proximately 2 million to approximately 11 thousand years be
fore the present time.
PLEURAL BONE — One of the elongate bones enclosing a rib and
making up the lateral part of the shell of a turtle.
PLIOCENE — A geologic epoch which extended from approxi
mately 12 million to approximately 2 million years before the
present time.
POLYTYPIC — A term used to describe a taxonomic category
containing more than one immediately subordinate category.
POSTFRONTAL — A bone which makes up the upper part of the
orbit in the turtle skull.
POSTORBITAL — A scale or other structure behind the eye.
POSTZYGAPOPHYSIS — One of two small projections extending
from the rear of a vertebra, which make sliding contact with
the prezygapophysis of the adjacent vertebra to the rear.
PRECENTRAL — Alternative name for the nuchal scute.
PRECOSTAL SCUTE — A scute preceding the costal scutes, pres
ent in a few extinct turtle species.
PREFRONTAL — A bone in the turtle skull which extends from
the nasal opening to the anterior part of the orbit.
PREMAXILLA — One of the two mesially contiguous bones which
extend from the nasal opening to the jaw and palate at the front
of a turtle skull.
PREZYGAPOPHYSIS — See postzygapophysis.
PROCOELOUS — A term used to describe a vertebra with an an
terior articular concavity.
881
www.Ebook777.com
PRONEURAL — Alternative name for the nuchal bone.
PTERYGOID — One of two mesially contiguous bones which
make up the rear part of the palate of a turtle.
PYGAL — The rearmost median bone in a turtle carapace.
QUADRATE — A bone toward the rear of a turtle shell which pro
vides the hinging surface for the lower jaw.
RADIUS — One of the two bones making up the forearm; in man
this bone is located on the same side as the thumb.
RAMUS — One branch or lateral extension of a symmetrical or
branched structure.
RATHKE’S GLANDS — A series of ventrolaterally situated
glands, present in many turtles, which in some species can
exude a highly odiferous secretion.
RETICULATIONS — A network of intercrossed lines.
SACRAL — Pertaining to the sacrum, or that part of the vertebral
column adjacent to the pelvis.
SCAPULA — A bone in the shoulder girdle with which the humer
us makes contact.
SEAM — The line of contact between two scutes.
SECONDARY PALATE — A posterior expansion of the alveolar
surfaces of certain turtles, particularly those adapted for a diet
of hard-shelled organisms.
SEPTUM — A thin, typically cartilaginous plate separating two
structures or vacuities.
SERRATED — Having a saw-toothed edge.
SPLENIAL — A bone located on the inside edge of the lower jaw,
toward the rear.
SQUAMOSAL — A bone located above the ear region of a turtle.
SULCUS — The impression left on the underlying bone by a seam
between scutes.
SUPRACAUDAL SCUTES — The two rearmost marginal scutes,
sometimes fused into one.
SUPRAMARGINAL SCUTES — Scutes present in Macroclemys,
located between the costals and the marginals.
SUPRAOCCIPITAL — The bone which forms the roof of the fora
men magnum; the posteriorly directed process formed by this
bone.
SUPRAPYGAL BONES — Those bones in the median series
which lie between the posteriormost neural bone and the pygal
bone.
SURANGULAR — A bone located near the rear of each ramus of
the lower jaw.
SUTURE — A jagged connection between two bones.
SYMPATRIC — Occupying the same geographic area.
SYMPHYSIS — A suture between two symmetrical structures, as
882
in the lower jaw of many animals or between the two halves of

the pelvis.
SYNONYM — One of several names which apply to the same taxo
nomic entity, only one of which is valid. In normal use, only the
invalid names are called synonyms.
TACHYCARDIA — The slowing of the heart that takes place when
a turtle dives.
TECTIFORM — Roof-shaped—having a high central ridge and
flat sides.
TEMPORAL — The sides of the skull behind the eyes.
TERTIARY — A geologic period encompassing Paleocene to Plio
cene epochs.
TIBIA — The thicker of the two parallel bones forming the lower
part of the hind leg.
TRICARINATE — Having three longitudinal ridges or keels.
TRITURATING — The functional surface of the jaw.
TUBERCLE — A small lump or knotlike projection.
TUBEROSITY - A large tubercle.
TYMPANUM — The vibrating surface which forms the external
ear in turtles.
TYPE — The specimen on which a species is based and named,
which should be kept available for permanent reference.
TYPE GENUS — The genus selected as a standard of reference
for a family, useful in cases where a later systematist decides
that a family should be split.
TYPE LOCALITY — The location from which the type specimen
or specimens of a species were collected.
TYPE SPECIES — The species upon which the description of a
genus is based. If a genus is later split into several genera, the
type species has to remain with the original generic designa
tion.
TYPICAL SUBSPECIES — That subspecies which bears the same
name as the species, e.g. Chrysemys picta picta.
ULNA — A bone in the forearm which runs parallel to the radius.
UNGULIGRADE — A gait in which only the claws make contact
with the ground.
VERMICULATIONS — Wavy or wormlike lines or patterns.
VERTEBRAL SCUTES — Alternative name for central scutes.
VOMER — An elongate median bone present in the palate of most
turtles.
WALLACE’S LINE — The imaginary boundary separating Asia
tic and Australian fauna; it passes between Bali and Lombok
and between Borneo and Celebes, east of the Philippines.
XERIC — A term used to describe dry or arid environments.
XIPHIPLASTRA — The rearmost pair of plastral bones.
883
www.Ebook777.com
ABOUT THE AUTHOR
Peter C. H. Pritchard was born in England in 1943, and was

resident in Belfast, Northern Ireland, from 1952 to 1961, where his
father is Professor of Anatomy at Queen’s University. He entered
Magdalen College, Oxford, in 1961, having been awarded an open
scholarship in chemistry. He graduated with Honours in 1964, in
which year he entered the University of Florida as a zoology
graduate student. He obtained his Ph.D. from that institution in
1969. From 1969 to 1973 Dr. Pritchard worked on marine turtle con
servation with the International Union for the Conservation of
Nature and the World Wildlife Fund.
His present position is Vice President for Science and
Research of the Florida Audubon Society, and he is also an
Adjunct Professor of Biology at Florida Technological University
and Florida Atlantic University. His field work took him to the
Guianas, especially French Guiana, every summer for almost a
decade, and he has undertaken six expeditions to the Galapagos
Islands. He has worked closely with the Mexican Government on
sea turtle conservation, and in 1976 he undertook a month-long
study of the marine turtles of Micronesia. He has also worked in
Iran, Peru, Ecuador, Honduras, and the Caribbean, and has
published about fifty papers on sea turtles as well as popular
articles on a variety of natural history subjects. Current projects
include the preparation of a monograph on the turtles of Venezuela
in cooperation with the Venezuelan National Parks Department.
Dr. Pritchard’s wife, Sibille, is the former Woman’s Editor of
the Guyana Graphic. They have two sons, Sebastian, born in 1972,
and Dominic, born in 1975.
885
INDEX
(BOLD numbers refer to color photos, italic numbers to other il

lustrations. )
abaxillare, Kinosternon, 540-541, 544 Batagur, baska, 194-196, 296, 558

abingdoni, Geochelone elephantopus, 344, 378-379, 382 bauri, Kinosternon bauri, 513, 5/6-517, 520
abrupta, Geochelone (fossil), 286 bauri, Terrapene carolim, 165, 167, 169, 280, 510
Acinixys planicauda, 361, 468-469 bealei, Sacalia, 190- 191. 293
acutirostris, Chelydra serpentina, 365, 488, 492 bechuanicus, Pelusios, 7 8
acutum, Kinosternon, 376, 545, 548 becki, Geochelone elephantopus, 333, 351, 354-355, 575
adansoni, Pelusios, 610, 763 belli, Chrysemys picta, 137, 503
Adanson’s turtle, 610, 763 belliana, Kinixys belliani, 458-459, 460-461, 582
Afghan tortoise, 349, 410-411, 414 Bell’s hinged tortoise, 458-459, 469461, 582
africana, Geochelone (fossil), 310 berlandieri, Gopherus, 422-423, 426, 587
agassizi, Chelonia mydas, 409, 684 bibroni, Pelochelys, 652 -653
agassizi, Gopherus, 352, 418-419, 422, 587 Big Bend mud turtle, 373, 540
Alabama map turtle, 139-142, 249, 252 Big Bend turtle, 112-//3
Alabama red-bellied turtle, 133-134, 248 Big-headed turtle, 401, 666-667, 668-669, 670
alabamensis, Pseudemys, 133-134, 248 Black pond turtle, 218-219, 320, 562
albertisii, Emydura australis, 806 Black-knobbed sawback, 142, 256
Aldabra giant tortoise, 287, 290-291, 294-295, 298-299, Black-rayed soft-shelled turtle, 396, 646-647
302, 325, 578 blandingi, Emydoidea, 157-158, 269, 531
Alligator snapping turtle, 368, 493, 496-497, 499, Blanding’s turtle, 157-158, 269, 531
500-501, 504 Bog turtle, 160, 161-162, 272, 535
Amboina box turtle, 211-213, 313, 551 Bolson tortoise, 431, 434435, 438, 586
amboinensis, Cuora, 211- 213, 313, 551 bomeensis, Orlitia, 197- 198, 297
Amphichelydia (fossil history), 74-78 bomeoensis, Callagur, 196- 197, 297
Amyda, see Trionyx boulengeri, Homopus, 449, 451-452, 453
angulata, Chersine, 357, 461-462, 463 Bowsprit tortoise, 357, 461-462, 463
angusta, Pseudemys decussata, 122-123 Box turtles, 163-172
angustatus, Claudius, 385, 580-581, 584-585, 588-589 Brahminy river turtle, 193-194, 558-559
angustipons, Kinosternon, 377, 487, 552 brasiliensis, Pseudemys scripta, 120-121
Annamemys annamensis, 191-192, 296 Bridled soft-shelled turtle, 388, 617, 620 -621
annamensis, Annamemys, 191 -192, 296 Burmese brown tortoise, 311, 314, 328, 582
annandalei, Hieremys, 36, 213, 214 -215 Burmese soft-shelled turtle, 646
annulata, Rhinoclemys, 178-179, 288 Burmese star tortoise, 274-215
Aperotemporalidae (fossil), 76-77 caglei, Graptemys, 149-150, 264
arachnoides, Pyxis, 251, 360, 466-468 ‘calcarata, * Geochelone, see G. sulcata
Archaeochelydium (fossil), 74 Callagur bomeoensis, 196-191, 297
Archelon (fossil), 96 callirostris, Pseudemys scripta, 119-120, 519
areolata, Rhinoclemys, 774-176, 547 Callopsis, see Rhinoclemys
areolatus, Homopus, 450, 452-453, 587 calvatus, Trionyx muticus, 393, 637
Arrau river turtle, 420, 482-483, 606, 631, 751-752 carajasensis, Kinosternon scorpioides, 529, 532
asper, Trionyx spinifer, 392, 598, 628-629, 632 Carapace osteology, 19, 20, 22, 24
ater, Trionyx, 625, 628 carbonaria, Geochelone, 326-327, 330-331, 332,
aubryi, Cycloderma, 617 570-571, 821
Aubry’s soft-shelled turtle, 617 Caretta caretta, 412, 590-591, 631, 693-695, 696-701
Australian side-necks, genera, 794 caretta, Caretta, 412, 590-591, 631, 693-695, 696-701
australis, Emydura australis, 611, 806 Carettochelyidae, 656-661
babcocki, Geochelone pardalis, 266- 267, 566 Carettochelyidae (fossil), 90-91
Baenidae (fossil), 76 Carettochelys insculpta, 400, 627, 630, 656, 657-658,
barbouri, Graptemys, 138-139, 249, 506 659-661
Barbour’s map turtle, 138-/39, 249, 506 carinatus, Pelusios, 766
baska, Batagur, 194-196, 296, 558 carinatus, Sternotherus, 380, 564-565
887
www.Ebook777.com
Carolina, Terrapene Carolina, 163-165, 280, 510, 638 Clemmys marmorata marmorata, 162-163, 273, 535
cartilagineus, Trionyx, 396, 646-647 Clemmys marmorata pallida, 162-163
Caspian turtle, 186, 757-188, 289, 539 Clemmys muhlenbergi, 160, 161- 162, 272, 535
caspica, Mauremys caspica, 186-187, 539 Coahuila box turtle, 168-170, 511
castaneus, Pelusios castaneus, 428, 766-767 coahuila, Terrapene, 168-170, 511
cataspila, Pseudemys scripta, 116-118, 232 Coastal plain turtle, 127, 128
cayannensis, Podocnemis, see P. erythrocephala Common batagur, 194-196, 296, 558
Central American river turtle, 401, 491, 661, Common box turtle, 163-165, 280, 510, 638
662-663, 664-665 Common map turtle, 145, 257
centrata, Malaclemys terrapin, 151-152 Common musk turtle, 380, 495, 560-561, 564
Chaco tortoise, 331, 332, 334-335, 574 Common snapping turtle, 33, 63, 364-365, 481,
chapini, Pelusios castaneus, 767 454-485, 488-489, 492-493, 498
chathamensis, Geochelone elephantopus, 344, 382-383, concinna, Pseudemys concinna, 127-128, 823
386, 578 coriacea, Dermochelys, 420, 594, 122-723, 724-727,
Chelidae, 771-812 728-732
Chelidae (fossil), 78-79 coronata, Melanochelys trijuga, 206-207
Chelodina, 795-803 Courtship, 61-62
Chelodina expansa, 444, 614, 799 crassicollis, Siebenrockiella, 224, 559
Chelodina longicollis, 445, 614-615, 795-796 creaseri, Kinosternon, 544-545
Chelodina novaeguineae, 796-797 cretica, mauremys caspica, 188
Chelodina oblonga, 119-800, 801 Crucilla turtle, 385, 577, 580
Chelodina parkeri, 802- 803 cruentatum, Kinosternon scorpioides, 372, 487, 533, 536
Chelodina rugosa, 802 Cryptodira (fossil history), 83-98
Chelodina siebenrocki, 801-802 Cumberland turtle, 112, 228
Chelodina steindachneri, 797-199 Cuora amboinensis, 211- 213, 313, 551
Chelonia, 676-693 Cuora flavomarginata, 213, 316
Chelonia depressa, 409, 690-693 Cuora galbinifrons, 2\3
Chelonia mydas, 404-405, 408-409, 590-591, 211-212, 313, 551
Cuora trifasciata,
616-680, 681-690 Cuora yunnanensis, 212
Chelonia mydas agassizi, 409, 684 Cyclanorbis elegans, 616-617
Cheloniidae, 672-722 Cyclanorbis senegalenis, 612- 613, 616
Cheloniidae (fossil), 94-95 Cyclemys dentata, 208-209, 312
Cheloniidae, genera, 675 Cyclemys mouhoti, 206, 208-209, 312
Chelus fimbriatus, 36, 429, 432, 618, 112-773, 774-775 Cyclemys tcheponensis, 209, 2/5-211
Chelycarapockidae (fossil), 91 Cycloderma aubryi, 617
Chelydra serpentina serpentina, 33, 63, 364, 481, Cycloderma frenatum, 388, 601, 617, 620- 621
484- 485, 488, 492-493, 498 dahli, Phrynops, 772, 791-192
Chelydra serpentina acutirostris, 365, 488-492 darwini, Geochelone elephantopus, 340, 366-367
Chelydra serpentina osceola, 364, 488-459, 492, 499 decorata, Pseudemys, 125- 126, 237
Chelydra serpentina rossignoni, 365, 488-489, 492 decussata, Pseudemys, 122-123
Chelydridae, 481-504 decussata, Pseudemys decussata, 122-123
Chelydridae (fossil), 83 Deirochelys, 154-157
Chersine angulata, 357, 461-462, 463 Deirochelys reticularia reticularia, 156, 268
chilensis, Geochelone chilensis, 331, 332, 334-335, 574 Deirochelys reticularia chrysea, 156, 268, 515
Chinemys grangeri, 221-222 Deirochelys reticularia miaria, 155- 156
Chinemys kwangtungensis, 222-223, 321 delticola, Graptemys niginoda, 142
Chinemys megalocephala, 223 dentata, Cyclemys, 208-209, 312
Chimneys reevesi, 219, 220-221, 321, 554 dentata, Elseya, 807, 808- 809
Chinese soft-shelled turtle, 397, 602, 641-649 denticulata, Geochelone, 323, 326-327, 331, 570-571
Chitra indica, 397, 650, 651- 652 depressa, Chelonia, 409, 690-693
chrysea, Deirochely reticularia, 156, 268, 515 depressa, Heosemys, 204
Chrysemys, 135-137 Depressed musk turtle, 384, 569, 572
Chrysemys picta picta, 135-136, 502 depressus, Sternotherus minor, 384, 569, 572
Chrysemys picta belli, 137, 503 derbianus, Pelusios castaneus, 161
Chrysemys picta dorsalis, 137, 502 Dermatemyidae, 661-665
Chrysemys picta marginata, 136, 248 Dermatemyidae (fossil), 84-85
Circulation, 54-59 Dermatemys mawi, 401, 491, 661, 662-663, 664-665
circumdata, Kachuga tentoria, 198-199, 300 Dermochelyidae, 722-732
Classifications, family, 98-103 Dermochelyidae (fossil), 98
Claudius angustatus, 385, 580-581, 584-585, 588-589 Dermochelys coriacea, 420, 594, 122-723, 724-727,
Clemmydopsis (fossil), 55 728-732
Clemmys, 158-163 Desert tortoise, 352, 418-419, 422, 587
Clemmys guttata, 159-160, 272, 534 dhongoka, Kachuga, 199- 200
Clemmys insculpta, 158-159, 269, 534 diademata, Rhinoclemys, 180, 182
888
Diamondback terrapin, 150-154, 264, 514 Florida box turtle, 165, 167, 169, 280, 510
Digestion, 53-54 Florida chicken turtle, 156, 268, 515
Dinner-plate turtle, 797 -799 Florida gopher tortoise, 20-21, 352, 426-427,
Dogania, see Trionyx subplanus 430-431, 586
donosobarrosi, Geochelone chilensis, 334-335 Florida mud turtle, 369, 509, 512
dorbignyi, Pseudemys scripta, 120- 121, 236, 530 Florida red-bellied turtle, 132-133, 245, 527
dorsalis , Chrysemys picta, 137, 502 Florida soft-shelled turtle, 24-25, 389, 598, 621,
dumeriliana, Podocnemis, see Peltocephalus tracaxa 624 -625
dunni, Kinosternon, 552-553, 556 floridana, Pseudemys floridana, 127, 128
Eastern chicken turtle, 156, 268 Fly River turtle, 400, 627, 630, 656, 657-658, 659-661
Eastern mud turtle, 491, 508-509 formosus, Trionyx, 646
Eastern painted turtle, 135-136, 502 forsteni, * Geochelone, 319
edeniana, Melanochelys trijuga, 205, 207 frenatum, Cycloderma, 388, 601, 617, 620 -621
Eggs, 65-70 funerea, Rhinoclemys, 177, 285, 542
Egyptian tortoise, 414-415, 418 Gabon turtle, 425, 761-762
elegans, Cyclanorbis, 616-617 gabonensis, Pelusios, 425, 761-762
elegans, Geochelone, 270-271, 274-275, 567 gadowi, Geochelone (fossil), 310
elegans, Pseudemys scripta, 112, 117, 225, 522-523 gaigeae, Pseudemys scripta, 112-113
elephantopus, Geochelone, 325, 335-394 galapagoensis, Geochelone elephantopus, 387, 390-391
elephantopus, Geochelone elephantopus, 337, 351, 359 Galapagos giant tortoises, 325, 335-394
362-363, 366, 574 galbinifrons, Cuora, 213
elongata, Geochelone, 315, 318, 329, 583 Ganges soft-shelled turtle, 599, 642 -643
Elseya, 807-810 gangeticus, Trionyx, 599, 642 -643
Elseya dentata, 807, 808 -809 Geochelone, 258-394
Elseya latisternum, 444, 809 Geochelone abrupta (fossil), 286
Elseya novaeguineae, 448, 809-810 Geochelone africana (fossil), 310
emoryi, Trionyx spinifer, 392, 628-629, 633, 636 Geochelone (Aldabrachelys), 286-302
Emydidae, 108-246 Geochelone (Asterochelys), 275-286
Emydidae (fossil), 85-86 Geochelone ‘calcarata, * see G. sulcata
Emydoidea blandingi, 157- 158, 269, 531 Geochelone carbonaria, 326-327, 330-331, 332,
Emydura, 803-807 570-571, 821
Emydura australis australis, 611, 806 Geochelone (Chelonoidisf 322-394
Emydura australis albertisii, 806 Geochelone chilensis chilensis, 331, 332, 334-335, 574
Emydura australis subglosbosa, 806-807 Geochelone chilensis donosobarrosi, 334-335
Emydura kreffti, 611, 805-806 Geochelone (Cylindraspis) (fossil), 302-310
Emydura macquarrii macquarrii, 803, 804 -805 Geochelone denticulata, 323, 326-327, 331, 570-571
Emydura macquarrii signata, 805 Geochelone elegans, 210-271, 274-275, 567
emys, Geochelone, 311, 314, 328, 582 Geochelone elephantopus, 325, 335-394
Emys orbicularis, 41, 56-57,183, 184-185, 289, 531 Geochelone elephantopus elephantopus, 337, 351, 359,
ephippium, Geochelone elephantopus, 341, 367, 370-371, 362-363, 366, 574
374, 579 Geochelone elephantopus abingdoni, 344, 378-319, 382
Eretmochelys imbricata, 416-417, 595, 673-674, 713-722 Geochelone elephantopus becki, 333, 351, 354-355, 575
erosa, Kinixys, 455-456, 457 Geochelone elephantopus chathamensis, 344, 382-383,
Erymnochelys madagascariensis, 424, 425, 760 386, 578
erythrocephala, Podocnemis , 606, 754-755 Geochelone elephantopus darwini, 340, 366-367
Eunotosaurus africanus (fossil), 73 Geochelone elephantopus ephippium, 341, 367, 370-371,
Euphrates soft-shelled turtle, 641 374, 579
euphraticus, Trionyx, 641 Geochelone elephantopus galapagoensis, 387, 390-391
European pond turtle, 41, 56-57, 183, 184-185, Geochelone elephantopus guntheri, 337, 351, 359,
289, 531 362-363, 366
expansa, Chelodina, 444, 614, 799 Geochelone elephantopus hoodensis, 340, 374-375, 378,
expansa, Podocnemis, 420, 482-483, 606, 631, 751-752 837
False map turtle, 145-147, 260 Geochelone elephantopus microphyes, 336, 351, 355, 358
felis, Pseudemys, 124, 530 Geochelone elephantopus phantastica, 345, 386-387
femoralis, Homopus, 452-453, 454 Geochelone elephantopus porteri, 333, 343, 346-347,
ferox, Trionyx, 24-25, 389, 598, 621, 624 -625 350, 579
fimbriatus, Chelus, 36, 429, 432, 618, 112-773, 774-775 Geochelone elephantopus vandenburghi, 336, 339, 351,
Flap-shelled turtle, 602, 608 600, <> I t 358-359
Flatback turtle, 409, 690-693 Geochelone elephantopus wallacei, 391, 394
Flat-headed flat-shelled turtle, 626, 778-779 Geochelone elongata, 315, 318, 329, 583
flavescens, Kinosternon flavescens, 494, 520-521, 524 Geochelone emys, 311, 314, 328, 582
flavimaculata, Graptemys, 142, 143, 144, 253, 506 Geochelone forsteni, ’ 319
flavomarginata, Cuora, 213, 316 Geochelone gadowi (fossil), 310
flavomarginatus, Gopherus, 431, 434 435, 438, 586 Geochelone (Geochelone), 259-275
889
www.Ebook777.com
Geochelone gigantea, 287, 290-291, 294-295, 298-299, Guau turtle, 384, 494, 573, 576-511
302, 325, 578 Gulf coast box turtle, 167-168, 281, 507
Geochelone gouffei (fossil), 287 guntheri, Geochelone elephantopus, 337, 351, 359,
Geochelone grandidieri (fossil), 286-287 362-363, 366
Geochelone hestema (fossil), 286-287 guttata, Clemmys, 159-160, 272, 534
Geochelone impressa, 314, 328 hamiltoni, Geoclemys, 218- 219, 320, 562
Geochelone indica (fossil), 302-303, 306-307 Hardella thurji, 193- 194, 558-559
Geochelone (IndotestudoX 314-319 hartwegi, Trionyx spinifer, 628-629, 632
Geochelone (ManouriaX 311-314 Hawksbill turtle, 416-417, 595, 673-674, 713-722

Geochelone pardalis pardalis, 263, 266-267, 270 Helmeted turtle, 428, 759, 779-771
Geochelone pardalis babcocki, 266- 267, 566 Heosemys depressa, 204
Geochelone platynota, 274- 275 Heosemys grandis, 109, 201-203, 305, 538, 550
Geochelone radiata, 275, 278-279, 282, 324, 567 Heosemys leytensis, 204
Geochelone sumeiri (fossil), 287 Heosemys spinosa, 202, 308
Geochelone sulcata, 259, 262-263, 324 Heosemys silvatica, 202-204
Geochelone travancorica, 318-319, 329, 583 hermanni, Testudo hermanni, 402-403, 406-407, 566
Geochelone vosmaeri (fossil), 307 Hermann’s tortoise, 348, 402-495, 406-407, 566
Geochelone yniphora, 279, 282-283, 286, 325 herrerai, Kinosternon, 376, 548-549, 552
Geoclemys hamiltoni, 218, 219, 320, 562 Herrera’s mud turtle, 376, 548-549, 552
Geoemyda spengleri spengleri, 201, 304 hestema, Geochelone (fossil), 330
Geoemyda spengleri japonica, 201 Hieremys annandalei, 36, 213, 274-215
geoffroanus, Phrynops geoffroanus, 437, 619, 622, Hieroglyphic turtle, 129, 518, 634
786- 787, 788- 789 hieroglyphica, Pseudemys concinna, 129, 518, 634
geographica, Graptemys, 145, 257 hilarii, Phrynops, 437, 186-787

Geometric tortoise, 353, 438-439 hiltoni, Pseudemys scripta, 114
geometricus, Psammobates, 353, 438-439 hippocrepis, Kinosternon subrubrum, 572-513
Giant musk turtles, 572-580 hirtipes, Kinosternon hirtipes, 373, 486, 537, 540
gibba, Phrynops, 432-433, 622, 782- 783 hogei, Phrynops, 785-786
gigantea, Geochelone, 287, 290-291, 294- 295, 298-299, homeana, Kinixys, 456-457, 458
302, 325, 578 Home’s hinged tortoise, 456-457, 458
Glyptopsidae (fossil), 77 Homopus, 449-454
Gopher tortoise, 20-21, 352, 426-427, 430-431, 586 Homopus areolatus, 450, 452-453, 587
Gopherus, 418-438 Homopus boulengeri, 449, 451-452, 453
Gopherus agassizi, 352, 418-419, 422, 587 Homopus femoralis, 452-453, 454
Gopherus berlandieri, 422-423, 426, 587 Homopus signatus, 449, 457
Gopherus flavomarginatus, 431, 434- 435, 438, 586 hoodensis, Geochelone elephantopus, 340, 374-375, 378,
Gopherus polyphemus, 20-21, 352, 426-427, 430-431, 586 837
gouffei, Geochelone (fossil), 287 horsfieldi, Testudo, 349, 410-477, 414
graeca, Testudo graeca, 395, 398-399, 402, 563 hoyi, Pseudemys floridana, 127-128, 240-241
grandidieri, Geochelone (fossil), 286-287 hurum, Trionyx, 393, 643-644
grandis, Heosemys, 109, 201-203, 305, 538, 550 Hydromedusa maximiliani, 115-776, 777-778
grangeri, Chinemys, 221-222 Hydromedusa tecdfera, 626, 115-776, 777-778
granosa, Lissemys punctata, 608, 613 ibera, Testudo graeca, 395, 398, 402
grand, Pseudemys, 126, 526 imbricata, Eretmochelys, 416-417, 595, 673-674, 713-722
Graptemys, 137-150 impressa, Geochelone, 314, 328

Graptemys barbouri, 138-139, 249, 506 Impressed tortoise, 314, 328
Graptemys caglei, 149-150, 264 incisa, Rhinoclemys pulcherrima, 175, 284, 543
Graptemys flavimaculata, 142, 143, 144, 253, 506 Indian roofed turtle, 198, 301, 555
Graptemys geographica, 145, 257 Indian star tortoise, 270-277, 274-275, 567
Graptemys kohni, 147- 148, 261, 503 indica, China, 397, 650, 957-652
Graptemys nigrinoda, 142, 256 indica, Geochelone (fossil), 302-303, 306-307
Graptemys oculifera, 143- 144, 253 indopeninsularis, Melanochelys trijuga, 205
Graptemys pseudogeographica pseudogeographica, insculpta, Carettochelys, 400, 627, 630, 656, 657-658,
145, 260 659-661

Graptemys pseudogeographica ouachitensis, 145, insculpta, Clemmys, 158-759, 269, 534
746-147, 260 integrum, Kinosternon, 536-537

Graptemys pseudogeographica sabinensis, 146-141 japonica, Geoemyda spengleri, 201
Graptemys pulchra, 139-142, 249, 252 japonica, Mauremys, 189
Graptemys versa, 148- 149, 261 Kachuga dhongoka, 199-200
grayi, Pseudemys scripta, 115- 116 Kachuga kachuga, 200
Green Turtle, 404-405, 408-409, 590-591, 616-680, kachuga, Kachuga, 200
681-690 Kachuga smithy , 199, 301, 555
Growth, shell, 29-32 Kachuga sylhetensis, 200-201

guadalupensis, Trionyx spinifer, 628-629, 633 Kachuga tecta, 198, 301, 555
890
Kachuga tentoria tentoria, 198-199 leytensis, Heosemys, 204

Kachuga tentoria circumdata, 198-199, 300 Lissemys punctata punctata, 602, 608, 613
Kachuga trivittata, 200 Lissemys punctata granosa, 608, 613
Karroo tortoise, 452-453, 454 Lissemys punctata scutata, 608-609, 613
kempi, Lepidochelys, 36, 413, 595, 701-707 littoralis, Malaclemys terrapin, 154
Kemp’s ridley, 36, 413, 595, 701-707 Loggerhead musk turtle, 381, 495, 565, 568-569
Keys, family, 9-10 Loggerhead turtle, 412, 590-591, 631, 693-695,
Keys, generic: 696-701
Chelidae, 11 longicollis, Chelodina, 445, 614-615, 195-796
Cheloniidae, 10 ‘lunata, ’ Rhinoclemys, 182
Chelydridae, 12 luteola, Terrapene omata, 170, 276
Emydidae, 14-16 lutescens, Pelusios williamsi, 768-769
Kinosternidae, 12 macquarrii, Emydura macquami, 803, 804-8 05
Pelomedusidae, 10-11 Macrobaenidae (fossil), 77-78
Testudinidae, 12-13 Macroclemys temmincki, 368, 493, 496-497, 499,
Trionychidae, 17 500-501, 504
Kinixys, 454-461 macrospilota, Malaclemys terrapin, 154, 265, 514
Kinixys belliana belliana, 458-459, 460-461, 582 madagascariensis, Erymnochelys, 424, 425, 760
Kinixys belliana nogueyi, 458, 460 major, Terrapene Carolina, 167-168, 281, 507
Kinixys erosa, 455-456, 457 Malaclemys, 150-154
Kinixys homeana, 456-457, 458 Malaclemys terrapin terrapin, 151-152
Kinosternidae, 504-589 Malaclemys terrapin centrata, 151-152
Kinosternidae (fossil), 83-84 Malaclemys terrapin littoralis, 154
Kinosternon, 505-557 Malaclemys terrapin macrospilota, 154, 265, 514
Kinosternon abaxillare, 540-541, 544 Malaclemys terrapin pileata, 154
Kinosternon acutum, 376, 545, 548 Malaclemys terrapin rhizophorarum, 152-153
Kinosternon angustipons, 377, 552, 487 Malaclemys terrapin tequesta, 152, 265
Kinosternon creasen, 544-545 Malacochersus tomieri, 26, 356, 463-465, 466
Kinosternon dunni, 522-553, 556 Malayan snail-eating turtle, 223-224, 562
Kinosternon flavescens flavescens, 494, 520-521, 524 Malayan soft-shelled turtle, 396, 647
Kinosternon flavescens spooneri, 520-521, 524 Malayemys subtrijuga, 223- 224, 562
Kinosternon flavescens stejnegeri, 520-521 malonei, Pseudemys, 124, 522
Kinosternon herrerai, 376, 548-549, 552 manni, Rhinoclemys pulcherrima, 175, 543
Kinosternon hirtipes hirtipes, 373, 486, 537, 540 marginata, Chrysemys picta, 136, 248
Kinosternon hirtipes murrayi, 373, 540 marginata, Testudo, 349, 407, 410, 843
Kinosternon integrum, 536-537 Margined tortoise, 349, 407, 410, 843
Kinosternon leucostomum, 486, 505, 525, 528-529 marmorata, Clemmys marmorata, 162-163, 273, 535
Kinosternon scorpiodes scorpiodes, 372, 486, 490, Matamata, 36, 429, 432, 618, 112-773, 774-775
529, 532-533 Mauremys, 185-190
Kinosternon scorpioides carajasensis, 529, 532 Mauremys caspica caspica, 186-187, 539
Kinosternon scorpioides cruentatum, 372, 487, 533, 536 Mauremys caspica cretica, 188
Kinosternon scorpioides pachyurum, 529, 532 Mauremys caspica rivulata, 186-188, 289
Kinosternon scorpioides seriei, 529, 532 Mauremys japonica, 189
Kinosternon sonoriense, 490, 524-525 Mauremys leprosa,185-186, 539
Kinosternon spurelli, 529 Mauremys mutica, 188- 189,292, 538
Kinosternon subrubrum subrubrum, 491, 508-509 mazvi, Dermatemys, 401, 491, 661, 662-663, 664-665
Kinosternon subrubrum hippocrepis, 512-513 maximiliani, Hydromedusa, 115-776, 111-118
Kinosternon subrubrum steindachneri, 369, 509, 512 megalocephala, Chinemys, 223
klauberi, Terrapene nelsoni, 171- 172, 277 megacephalum, Platystemon megacephalum, 401,
kleinmanni, Testudo, 414-415 418 666-667, 668, 670
kohni, Graptemys, 147-148, 261, 503 Meiolania (fossil), 78
kreffti, Emydura, 611, 805-806 Meiolaniidae (fossil), 78
kwangtungensis, Chinemys, 222-223, 321 Melanochelys tricarinata, 204-205
latistenum, Elseya, 444, 809 Melanochelys trijuga trijuga, 205, 207, 309
Leatherback turtle, 420, 594, 122-723, 724-727, 728-732 Melanochelys trijuga coronata, 206-207
leithi, Trionyx, 643 Melanochelys trijuga edeniana, 205, 207
Leopard tortoise, 263, 266-267, 270, 566 Melanochelys trijuga indopeninsularis, 205
Leopard tortoise, 263, 266-267, 270, 566 Melanochelys trijuga parkeri, 205
Lepidochelys, 701-713 Melanochelys trijuga thermalis, 206, 309, 550
Lepidochelys kempi, 36, 413, 595, 701-707 melanosterna, Rhinoclemys, 180-181, 288
Lepidochelys olivacea, 416, 594, 101-709, 710-713 Mexican red turtle, 173-175, 284, 543
leprosa, Mauremys, 185-186, 539 mexicana, Terrapene Carolina, 16, 168, 281
leucostomum, Kinosternon, 486, 525, 528-529 miaria, Deirochelys reticularia, 155- 156
lezuyana, Podocnemis, 607, 155-756, 757 microphyes, Geochelone elephantopus, 336, 351, 355, 358
891
www.Ebook777.com
Midland painted turtle, 136, 248 vertebral column, 35-40
minor•, Stemotherus minor, 381, 495, 565, 568-569 Ouachita map turtle, 145, 746-147, 260
Mississippi diamondback terrapin, 154 ouachitensis, Graptemys pseudogeographica, 145,
Mississippi map turtle, 147-148, 261, 503 746-147, 260

Mississippi mud turtle, 512-513 pachyurum, Kinosternon scorpio ides, 529, 532
Missouri slider, 127-128, 240-241 Pacific pond turtle, 162-163, 273, 535
Mobile turtle, 129-130, 244 pallida, Clemmys marmorata, 162-163
mobiliensis, Pseudemys concinna, 129-130, 244 pallidipectoris, Platemys, 119-780
Moorish tortoise, 395, 398-399, 402, 563 pallidus, Trionyx spinifer,628-629, 632-633
Morenia ocellata, 217 palmarum, Kinosternon bauri, 576-517
Morenia petersi,217-218 Pancake tortoise, 26, 356, 463-465, 466
mouhoti, Cyclemys, 206, 208-209, 312 pardalis, Geochelone pardalis, 263, 266-267, 270
Mud turtles, 505-557 parkeri, Chelodina, 802- 803
muhlenbergi, Clemmys, 160, 161- 162, 272, 535 parkeri, Melanochelys trijuga, 205
Murray River turtle, 803, 804-805 Parrot-beaked tortoise, 450, 452-453, 587
murrayi, Kinosternon hirtipes, 373, 540 Peacock soft-shelled turtle, 393, 643-644
Muscles, 44-50 peguense, Platystemon megacephalum, 666, 668-669
Musk turtles, 557-572 Pelochelys bibroni, 652- 653

mutica, Mauremys, 188- 189, 292, 538 Pelomedusa subrufa, 428, 756, 770-11 1
muticus, Trionyx muticus, 636-637, 640 Pelomedusidae, 749-771

mydas, Chelonia, 404-405, 408-409, 590-591, 676-680, Pelomedusidae (fossil), 79-82
681-690 Pelomedusidae, genera, 749-750
Nagpur soft-shelled turtle, 643 peltifer, Stemotherus minor, 381, 569
Nanhsiungchelyidae (fossil), 91 Peltocephalus tracaxa, 158-759, 760, 822
nanus, Pelusios, 765 Pelusios, 761-769
Narrow-bridged mud turtle, 385, 580-581, 584-585, Pelusios adansoni, 610, 763
588-589 Pelusios bechuanicus, 768

Narrow-headed soft-shelled turtle, 397, 650, 651-652 Pelusios carinatus, 766
nasuta, Phrynops nasuta, 440, 623, 789-790 Pelusios castaneus castaneus, 428, 766-767
nasuta, Rhinoclemys, 179 Pelusios castaneus chapini, 767

nebulosa, Pseudemys scripta, 113-114, 229 Pelusios castaneus derbianus, 767
nelsoni, Pseudemys, 132-133, 245, 527 Pelusios castaneus rhodesianus, 767
nelsoni, Terrapene nelsoni, 171-172 Pelusios gabonensis, 425, 761-762
Nervous system, 59-61 Pelusios nanus, 765

New Guinea snapper, 448, 809-810 Pelusios niger, 764, 765
niger, Pelusios, 764, 765 Pelusios sinuatus, 762-163
nigricans, Trionyx, 644, 645- 646 Pelusios subniger, 607, 765-766
nigrinoda, Graptemys nigrinoda, 142 Pelusios williamsi williamsi, 768-769
Nile soft-shelled turtle, 599, 640-641 Pelusios williamsi lutescens, 168-769
nogueyi, Kinixys belliana, 458, 460 Peninsula turtle, 127, 241, 523
Notochelys platynota, 215- 216, 316 peninsularis, Pseudemys floridana, 127, 241, 523
novaeguineae, Chelodina, 796-797 perixantha, Rhinochelmys rubida, 176, 285
novaeguineae, Elseya, 448, 809-810 petersi, Morenia, 217-218
Nubian soft-shelled turtle, 616-617 phantastica, Geochelone elephantopus, 345, 386-387
oblonga, Chelodina, 799-800, 801 Phrynops (Batrachemysf 783-784, 789-794
Ocadia sinensis, 216-217, 317 Phrynops dahli, 772, 791-192
ocellata, Morenia, 217 Phrynops geoffroanus geo/froanus, 437, 619, 622,

oculifer, Psammobates, 353, 439, 442-443 786-181, 788-189
oculifera, Graptemys, 143- 144, 253 Phrynops geoffroanus tuberosus, 181-788,789

odoratus, Stemotherus, 380, 495, 560- 561, 564 Phrynops gibba, 432-433, 622, 782-183
oltvacea, Lepidochelys, 416, 594, 707-709, 710-713 Phrynops hilarii, 437, 186-787
Olive ridley, 416, 494, 101-709, 710-713 Phrynops hogei, 785-786
orbicularis, Emys, 41, 56-57, 183, 794-185, 289, 531 Phrynops (Mesoclemmysf 782-783
Orlitia borneensis, 797-198, 297 Phrynops nasuta nasuta, 440, 623, 189-790
omata, Pseudemys scripta, 114-115, 515 Phrynops nasuta wermuthi, 441, 623, 790-791
ornata, Terrapene omata, 769-170, 277 Phrynops (Phrynops), 783-789
Ornate box turtle, 769-170, 277 Phrynops rufipes, 436, 784-785
Ornate diamondback terrapin, 154, 265 Phrynops tuberculata, 792-193
osceola, Chelydra serpentina, 364, 488-489, 492, 499 Phrynops vanderhaegei, 793-194
Osteology: picta, Chrysemys picta,135-136, 502

hyoids, 44 154
pileata, Malaclemys terrapin,
limbs, 40-44 plana, Pseudemys decussata, 122-123
shell, 19-26 planicauda, Acinixys, 361, 468469
skull, 32-35 Plastron osteology, 21, 22, 25
892
Platemys pallidipectoris, 119-780 Pseudemys scripta elegans, 112, 117, 225, 522-523
Platemys platycephala, 626, 118-779 Pseudemys scripta gaigeae, 112-7 76
Platemys radiolata radio lata, 180-781 Pseudemys scripta grayi, 775-116
Platemys radiolata spixi, 180-781 Pseudemys scripta hiltoni, 114
platycephala , Platemys, 626 118-779 Pseudemys scripta nebulosa, 113-114, 229
platynota, Geochelone, 274-215 Pseudemys scripta ornata, 114-115, 515
platynota, Notochelys, 215- 216, 316 Pseudemys scripta taylori, 118
Platysternidae, 666-670 Pseudemys scripta troosti, 112, 228

Platysternidae (fossil), 83 Pseudemys scripta venusta, 118-119, 233
Platystemon megacephalum megaceph a ^ u m > 666-667, Pseudemys scripta yaquia, 114
668, 670 Pseudemys stejnegeri, 126

Platystemon megacephalum peguense, 666, 668-669 Pseudemys terrapen, 123-124, 237
Platystemon megacephalum vogeli, 668, 670 Pseudemys ‘vicina’, 125-126
Pleurodira (fossil history), 78-82 pseudogeographica, Graptemys pseudogeographica,
Pleurosternidae (fossil), 75-76 145, 260
Podocnemis, 750-758 Pseudotestudo, see Testudo kleinmanni
Podocnemis cayennensis, see P. erythr° ce phala Ptycogastridae (fossil), 86
Podocnemis dumeriliana, see Peltocephalus traeaxa pulcherrima, Rhinoclemys pulcherrima, 173
Podocnemis erythrocephala, 606, 754-755 pulchra, Graptemys, 139-142, 249, 252
Podocnemis expansa, 420, 482-483, 606, *31, 751-752 punctata, Lissemys punctata, 602, 608, 613
Podocnemis lewyana, 607, 155-756, 757 punctularia, Rhinoclemys, 179-180, 181, 182-183, 546
Podocnemis sextuherculata, 424, 755-756 putnami, Terrapene Carolina (fossil), 167
Podocnemis unifilis, 421, 603, 152-183, 754 Pyxidea, see Cyclemys
Podocnemis vogli, 757-158 Pyxis arachnoides, 251, 360, 466-468
polyphemus, Gopherus, 20-21, 352, 426 4 27, quadriocellata, Sacalia, 190-707
430-431, 586 radiata, Geochelone, 275, 218-279, 282, 324, 567
porteri, Geochelone elephantopus, 333* 343, 346-347, Radiated tortoise, 275, 218-279, 282, 324, 567
350, 579 radiolata, Platemys radiolata, 780-707
Proganochelyidae (fossil), 74-75 Razorback musk turtle, 380, 564-565

Proterochersidae (fossil), 75 Red-bellied turtle, 131-762, 245
Protostega (fossil), 97 Red-cheeked mud turtle, 372, 487, 533, 536
Protostegidae (fossil), 95-98 Red-eared turtle, 112, 117, 225, 522-523
Psammobates, 438447 Red-foot tortoise, 326-327, 330-331, 332, 570-571, 821
Psammobates geometricus, 353, 438-439 Reeves’ turtle, 219, 220-227, 321, 554
Psammobates oculifer, 353 , 439, 442^4 3 reevesi, Chinemys, 219, 220-221, 321, 554
Psammobates tentorius tentorius, 443* 347 Reproduction, 61-70
Psammobates tentorius triment, 443, 446-447 Reproductive organs, 61-63
Psammobates tentorius verroxii, 356, 447 Respiration, 50-53
Pseudemydura umbrina, 448, 810, 6/7-672 reticularia, Deirochelys reticularia, 156, 268
Pseudemys, 110-135 Rhinoclemmys, see Rhinoclemys
Pseudemys alabamensis, 133-134, 248 Rhinoclemys, 172-183
Pseudemys concinna concinna, 128-12^* 623 Rhinoclemys annulata, 178-179, 288

Pseudemys concinna hitroglyphica , l'29> 518, 634 Rhinoclemys areolata, 7 74-176, 547
Pseudemys concinna mobihensis, 129-1 30, 244 Rhinoclemys diademata, 180, 182
Pseudemys concinna sutoanniensit, 130* 527 Rhinoclemys funerea, 177, 285, 542
Pseudemys concinna texana, 130-/6/ Rhinoclemys lunata, ’ 182
Pseudemys decorata, 725-126, 237 Rhinoclemys melanosterna, 180-181, 288
Pseudemys decussata, 122-726 Rhinoclemys nasuta, 179

Pseudemys decussata angusta, 122-123 Rhinoclemys pulcherrima pulcherrima, 173
Pseudemys decussata plana, 122-123 Rhinoclemys pulcherrima incisa, 175, 284, 543
Pseudemys felis, 124, 530 Rhinoclemys pulcherrima manni, 175, 543
Pseudemys floridana floruiana, 127, 1^8 Rhinoclemys pulcherrima rogerbarbouri, 173-174
Pseudemys flondana hoyi, 127-128, 240-241 Rhinoclemys punctularia, 179-180, 181, 182-183,546
Pseudemys flondana peninsula ns, 127* 241, 523 Rhinoclemys rubida rubida, 176, 547
Pseudemys granti, 126, 526 Rhinoclemys rubida perixantha, 176, 285
Pseudemys inflata (fossil), 111 rhueophorarum, Malaclemys terrapin, 152-153
Pseudemys malonei, 124, 522 rhodestanus, Pelusios castaneus, 767
Pseudemys nelsoni, 132-1 33, 245, 527 Ridley turtle, 701-713
Pseudemys rubriventns, 131-762, 245 Ringed sawback, 746-144, 253
Pseudemys scripta scripta, 111, 51H River cooler, 128-129, 823
Pseudemys scripta bras 11 lent is, 120-121 nvulata, Mauremys caspica, 186-188, 289
Pseudemys sertpta calhrostns, 119 120< 519 robertmertensi, Testudo hermanm, 348, 403, 406
Pseudemys scripta cataspila, 116-118, 232 rogerbarbouri, Rhinoclemys pulcherrima, 173-174
Pseudemys scripta dorbignyi, 760-121, 236, 530 rossignom, Chelydra serpentina, 365, 488-489, 492
893
www.Ebook777.com
rubida, Rhinoclemys rubida, 176, 547 steindachneri, Kinosternon subrubrum, 369, 509, 512
rubriventris, Pseudemys, 131-132, 245 steindachneri, Trionyx, 649-650
rufipes, Phrynops, 436, 784-785 stejnegeri, Kinosternon flavescens,520-521
rugosa, Chelodina, 802 stejnegeri, Pseudemys, 126
Sabine map turtle, 746-147 Stemotherus, 557-572
sabinensis , Graptemys pseudogeographica, 746-147 Stemotherus carinatus, 380, 564-565
Sacalia bealei, 190-191, 293 Stemotherus minor minor, 381, 495, 565, 568-569
Sacalia quadriocellata, 190-191 Stemotherus minor depressus, 384, 569, 572
Salt Basin box turtle, 170, 276 Stemotherus minor peltifer, 381, 569
salvini, Staurotypus, 385, 577, 580 Stemotherus odoratus, 380, 495, 560- 561, 564
Saw-shelled snapper, 444, 809 Stinkpot, 380, 495, 560-561, 564
Schweigger’s hinge-back tortoise, 455-456, 457 Stripe-necked musk turtle, 381, 569
scorpioides, Kinosternon scorpioides, 372, 486, 490, 529, Striped mud turtle, 513, 516-517, 520
532- 533 subglobosa, Emydura australis, 806-807
Scorpion mud turtle, 372, 486, 490, 529, 532-533, 536 subniger, Pelusios, 607, 756-766
scripta, Pseudemys scripta, 111, 518 subplanus, Trionxy, 396, 647
scutata, Lissemys punctata, 608-609, 613 subrubrum, Kinosternon subrubrum, 491, 508-509
Scutes, 27-29 subrufa, Pelomedusa, 428, 750, 770-111

Sea turtles, 672-732 subtrijuga, Malayemys, 223- 224, 562
Senegal soft-shelled turtle, 672-613, 616 sulcata, Geochelone, 259, 262-263, 324
senegalensis, Cyclanorbis, 612-613, 616 sumeirei, Geochelone (fossil), 287
seriei, Kinosternon scorpioides, 529, 532 Suwannee turtle, 130, 527
serpentina, Chelydra serpentina, 33, 63, 364, 481, suwanniensis, Pseudemys concinna, 130, 527
484- 485, 488, 492-493, 498 swinhoei, Trionyx, 649
Serrated tortoise, 353, 439, 442-443 sylhetensis, Kachuga, 200-201
Serrated turtle, 762-763 taylori, Pseudemys scripta, 118
Sex ratios, 64-65 tcheponensis, Cyclemys, 209, 276-211
sextuberculata, Podocnemis, 424, 755-756 tecta, Kachuga, 198, 301, 555
Side-necked turtles, 748-812 tectifera, Hydromedusa, 626, 775-776, 777-778
siebenrocki, Chelodina, 801-802 temmincki, Macroclemys, 368, 493, 496-497, 499,
Siebenrockiella crassicollis, 224, 559 500-501, 504
signata, Emydura macquarrii, 805 Tent tortoise, 356, 443, 446-447
signatus, Homopus, 449, 451 tentoria, Kachuga tentoria, 198-199
silvatica, Heosemys, 202-204 tentorius, Psammobates tentorius, 443-447
Sinemydidae (fossil), 85 terrapen, Pseudemys, 123-124, 237

sinensis, Ocadia, 216-217, 317 Terrapene, 163-172
sinensis, Trionyx sinensis, 397, 648-649 Terrapene Carolina Carolina, 31, 69, 163-165, 280,
sinuatus, Pelusios, 762-763 510, 638
Sliders, 108-246 Terrapene Carolina bauri, 165, 167,7 69, 280, 510
smithi, Kachuga, 199, 301, 555 Terrapene Carolina major, 167-168, 281, 507
Smooth soft-shelled turtle, 393, 636-637, 640 Terrapene Carolina mexicana, 166, 168, 281
Snapping turtles, 481-504 Terrapene Carolina putnami (fossil), 167
Soft-shelled turtles, 600-655 Terrapene Carolina triunguis, 167, 507, 511
Soft-shelled turtles, genera, 605 Terrapene Carolina yucatana, 166, 168
Sonoran mud turtle, 490, 524-525 Terrapene coahuila, 168-170, 511
sonoriense, Kinosternon, 490, 524-525 Terrapene nelsoni nelsoni, 171-172
Southern painted turtle, 137, 502 Terrapene nelsoni klauberi, 777-172, 277
Spanish turtle, 185-766, 539 Terrapene omata omata, 766-170, 277
Speckled tortoise, 449-457 Terrapene omata luteola, 170, 276
spengleri, Geoemyda spengleri, 201, 304 terrapin, Malaclemys, 151-152, 264, 514
Spider tortoise, 257, 360, 466-468 tequesta, Malaclemys terrapin, 152, 265
spinifer, Trionyx, 392, 628-629, 632-633, 636 terrapin, Malaclemys terrapin, 151-152
spinifer, Trionyx spinifer, 628-629, 632, 636 terrestris, Testudo graeca, 398-399, 402
spinosa, Heosemys, 202, 308 Testudinidae, 247-480
Spiny soft-shelled turtle, 628-629, 632-633, 636 Testudinidae (fossil), 86-89
spixi, Platemys radiolata, 780-767 Testudo, 394-418
spooneri, Kinosternon flavescens, 520-527, 524 Testudo graeca graeca, 395, 398-399, 402, 563
Spotted turtle, 159-160, 272, 534 Testudo graeca ibera, 395, 398, 402
Spurred tortoise, 259, 262-263, 324 Testudo graeca terrestris, 398-399, 402
spurrelli, Kinosternon, 529 Testudo graeca zarudnyi, 398, 402, 563
Staurotypus, 572-580 Testudo hermanni hermanni, 402-403, 406407, 566
Staurotypus salvini, 385, 577, 580 Testudo hermanni robertmertensi, 348, 403, 406
Staurotypus triporcatus, 384, 494, 573, 576-511 Testudo horsfieldi, 349, 410411, 414
steindachneri, Chelodina, 797-199 Testudo kleinmanni, 414-415, 418
894
Testudo marginata, 349, 407, 410, 843 Trionyx spinifer guadalupensis, 628-629, 633
texana, Pseudemys concinna, 130-131 Trionyx spinifer hartwegi, 628-629, 632
Texas diamondback terrapin, 154 Trionyx spinifer pallidus, 628-629, 652-633
Texas map turtle, 745-149, 261 Trionyx steindachneri, 649-650
Texas slider, 130-757 Trionyx subplanus, 396, 647
Texas tortoise, 422-425, 426, 587 Trionyx swinhoei, 649
Thalassemydidae (fossil), 76 Trionyx triunguis, 599, 640-641
thermalis, Melanochelys trijuga, 206, 309, 550 triporcatus, Staurotypus, 384, 494, 573, 576-511
Three-toed box turtle, 167, 507, 511 triunguis, Terrapene Carolina, 167, 507, 511
thurji, Hardella, 795-194, 558-559 triunguis, Trionyx, 599, 640-641
Toad-headed turtle, 440-441, 623, 789-791 trivittata, Kachuga, 200
tomieri, Malacochersus, 26, 356, 463-465, 466 troosti, Pseudemys scripta, 112, 228
Tortoises, 247-480 tuberculata, Phrynops, 792-193
Toxochelidae (fossil), 91-94 602, 648-649
tuberculatus, Trionyx sinensis,
tracaxa, Peltocephalus, 758-759, 760, 822 tuberosus, Phrynops geoffroams, 181-788, 789
Travancore tortoise, 318-319, 329, 583 umbrina, Pseudemydura, 448, 810, 811-812
travancorica, Geochelone, 318-319, 329, 583 unifilis, Podocnemis, 421, 603, 152-753, 754
Triassochelys (fossil), 74-75 vandenburghi, Geochelone elephantopus, 336, 339, 351,
tricarinata, Melanochelys, 204-205 358-359
trijasciata, Cuora, 211-212, 313, 551 vanderhaegei, Phrynops, 795-794
trijuga, Melanochelys trijuga, 205, 207, 309 venusta, Pseudemys scripta, 118-119, 233
trimeni, Psammobates tentorius, 443, 446 -447 verroxii, Psammobates, tentorius, 356, 447
Trionychidae, 600-655 versa, Graptemys, 746,-149, 261
Trionychidae (fossil), 89-90 'vicina' Pseudemys, 125-126
Trionychidae, genera, 605 vogeli, Platystemon megacephalum, 668, 670
Trionyx, 605, 621-650 vogli, Podocnemis, 757-758
Trionyx ater, 625, 628 vosmaeri, Geochelone (fossil), 307
Trionyx cartilagineus, 396, 646-647 wallacei, Geochelone elephantopus, 391, 594
Trionyx euphraticus, 641 wermuthi, Phrynops nasuta, 441, 623, 790-791
Trionyx ferox, 24-25, 389, 598, 621, Western chicken turtle, 755-156
624 -625 Western painted turtle, 137, 503
Trionyx formosus, 646 White-lipped mud turtle, 486, 525, 528-529
Trionyx gangeticus, 599, 642 -643 williamsi, Pelusios williamsi, 768-769
Trionyx hurum, 393, 643-644 Wood turtle, 158-759, 269, 534
Trionyx leithi, 643 yaquia, Pseudemys scripta, 114
Trionyx muticus muticus, 636-637, 640 Yellow mud turtle, 494, 520-527, 524
Trionyx muticus calvatus, 393, 637 Yellow-bellied turtle, 111, 518
Trionyx nigricans, 644, 645 -646 Yellow-blotched sawback, 142, 143, 144, 253, 506
Trionyx sinensis sinensis, 397, 648-649 Yellow-foot tortoise, 323, 326-527, 331, 570-571
Trionyx sinensis tuberculatus, 602, 648-649 yniphora, Geochelone, 279, 282-283, 286, 325
Trionyx spinifer, 628-629, 632-633, 636 yucatana, Terrapene Carolina, 166, 168
Trionyx spinifer spinifer, 628-629, 632, 636 yunnanensis, Cuora, 212
Trionyx spinifer asper, 392, 598, 628-629, 632 Zambezi soft-shelled turtle, 388, 601, 617, 620 -621
Trionyx spinifer emoryi, 392, 626, 628-629, 633 zarudnyi, Testudo graeca, 398, 402, 563
895
www.Ebook777.com

0876669186

Enviado por

Dados do documento

Direitos autorais

Formatos disponíveis

Compartilhar este documento

Compartilhar ou incorporar documento

Opções de compartilhamento

Você considera este documento útil?

Este conteúdo é inapropriado?

Direitos autorais:

Formatos disponíveis

0876669186

Enviado por

Direitos autorais:

Formatos disponíveis

Free ebooks ==> www.Ebook777.

© 1979 T.F.H. Publications, Inc. Ltd.

Distributed in the U.S. by T.F.H. Publications, Inc., 211 West

KEY TO TURTLE FAMILIES

KEYS TO TURTLE GENERA

KEY TO THE CHELONIIDAE

KEY TO THE PELOMEDUSIDAE

KEY TO THE CHELIDAE

KEY TO THE KINOSTERNIDAE

KEY TO THE CHELYDRIDAE

KEY TO THE TESTUDINIDAE

1' Front lobe of plastron not moveable ...................................... 2

ridged; head and neck with longitudinal light stripes...........

KEY TO THE TRIONYCHIDAE

and plastron meet at a ‘bridge’ on each side. Both carapace and

OSTEOLOGY OF THE CARAPACE

OSTEOLOGY OF THE PLASTRON

VARIANT SHELL STRUCTURES

All living turtle species normally have eight pairs of pleural

Generalized scutes of the carapace (left) and plastron (right). Carapace: N

VARIANT SCUTE CONDITIONS

bryos) and phylogenetic studies (of fossils). However, there is no

SHELL GROWTH AND PIGMENTATION

Skull of the common snapping turtle, Chelydra serpentina , illustrating the

That part of the bodenaponeurosis that passes over the otic

CERVICAL AND DORSAL VERTEBRAE

usual among vertebrates. Moreover the lungs of turtles may be

produce a slight negative pressure in the lungs. Conversely, a tur­

The brachiocephalic artery divides into the two subclavian ar­

A pair of spinal nerves develops from each vertebra of the

Penis of Chelydra serpentina rossignoni.

Staurotypus is a relatively recently and independently derived one

Turtle eggs do not vary in size nearly as much as do the adult

Auffenberg, W. 1966. The carpus of land tortoises (Testudininae).

The Order Testudines has a long and distinguished history, and

Eunotosaurus africanus, once considered an ancestor to the turtles.

region. Because of the superficial similarity to the skull of the liv­

parallels that of the Meiolaniidae. On the other hand, the Pelome-

fajumensis from the Oligocene of Egypt, P. stromeri from the Eo­

nemys from the Eocene of Europe, and Peltochelys from the

Another fossil emydid genus was Broilia, from the Oligocene

tortoises, the alternating octagonal and quadrilateral neural

chilensis. The newly-described species G. hestema, from the late

and marginal scutes were defined by almost complete sulci, but

* European and African toxochelids are now known.

of the hyoplastron. Only a single species, Osteopygis emarginatus,

presence in only the former of intercostal fontanelles. Procolpo-

* Cimochelys benstedi, from the Upper Cretaceous of England,

Carapace and plastron of a gigantic extinct protostegid turtle, Archelon

The more primitive protostegids of the genera Chelosphargis

(I) Mlynarski, 1969

(III) Romer, 1956

(VI) Gaffney, 1975

Infraorder Pleurodira (Cope, 1868)

Dacque, E. 1912. Die fossilen Schildkroten Aegyptens. Geol. Palae-

Zangerl, R. 1945. Fossil specimens of Macrochelya from Tertiary

The family Emydidae is a large and varied assemblage of

and Webb in their description of Pseudemys scripta yaquia, it ap­

Pseudemys scripta grayi of the Pacific drainage of southern Mexico to El

Specimen of Pseudemys scripta elegans from Ciudad Victoria, Mexico, the

to have branching tails, and to be located near the middle of the

Pseudemys scripta dorbignyi from Argentina. If recognized as a full species,

introduced should be investigated, dorbignyi being sold in the pet

A uniform characteristic of the island turtles, shared by some

produce a slight negative pressure in the lungs. Conversely, a tur

The brachiocephalic artery divides into the two subclavian ar

region. Because of the superficial similarity to the skull of the liv

fajumensis from the Oligocene of Egypt, P. stromeri from the Eo

and Webb in their description of Pseudemys scripta yaquia, it ap

Along the Gulf Coast from the panhandle of Florida to Louisi

reticulations, and the entire surface is sculptured with fine longitu

The principal home of muhlenbergi is a block of land extend

of narrow radiating yellow streaks on a dark brown or black back

Hardella thurji, a common species of India and Pakistan. Photo by G. Mar

Young specimens of Cyclemys dentata are sometimes con

Specimen agreeing well with the description of Cyclemys tcheponensis, ten