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The
BIOLOGY
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SEA TURTLES
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The
BIOLOGY
of
SEA TURTLES
Edited by
Peter L. Lutz
John A. Musick
CRC PR E SS
Boca Raton London New York Washington, D.C.
Library of Congress Cataloging-in-Publication Data
The biology of sea turtles / edited by Peter L. Lutz and John A.

Musick.
p. cm. — (CRC marine science series)
Includes bibliographical references (p. ) and index.
ISBN 0-8493-8422-2
1. Sea turtles. I. Lutz, Peter L. II. Musick, John A.
III. Series; Marine science series.
QL666.C536B56 1996
597.92—dc20 96-36432
CIP
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Preface
The impetus for this book comes from the explosion in interest and involvement in
sea turtle biology and conservation. The interest in sea turtles is easy to understand.
The turtles belong to the most ancient line of living reptiles, first appearing more
than 200 million years ago in the late Triassic. When turtles first entered the sea is
not known, probably in the early Mesozoic, and for the next 100 million years,
during the rise and reign of the dinosaurs, the sea turtles shared the ocean with a
rich diversity of other air breathing reptiles, including the ichthyosaurs and plesio
saurs. But while the end of the Cretaceous witnessed the extinction of the dominant
large reptiles, the sea turtles continued to flourish up until very recent times when
their numbers have, through human interference, drastically declined. In order to
appreciate the causes of this collapse it must help to understand the selective advan
tages that have allowed sea turtles to survive so tenaciously, and, more particularly,
how they managed to hold their own against the comparatively recent invasions of
the sea by mammals. Up until the past 20 years or so most information on sea turtles
had been confined by the accessible, the nesting female and her eggs. But the use
of new technologies in data gathering — including mitochondrial DNA analyses,
remote sensing, and more sophisticated physiological monitoring techniques — has
resulted in a substantially greater understanding of sea turtle biology at all stages
of their life history. Major advances have been made in such broad areas as popu
lation genetics and phylogeny, sensory biology, migration and orientation, hatchling
behavior, age and growth, reproduction and endocrinology, sex determination, diving
physiology, and osmoregulation.
The results of these diverse studies are widely scattered in the literature which
this book brings together in a comprehensive summary; no similar reference has
previously been available. However, it is not a mere compilation, its focus is on how
sea turtles operate in, are adapted to, and are dependent upon their marine environ
ment. Considering the endangered status of these fascinating giant marine reptiles,
the book also deals with the threats to survival presented by manmade changes to
the ocean and coastal zones.
Peter L. Lutz
John A. Musîck
Acknowledgment
The idea for this book was bom out of many late nights of discussions with friends
and colleagues at annual meetings of the Sea Turtle Symposium. This convocation
has grown since 1981 from a small group of about 60 scientists and conservationists
to more than 700 workers drawn from 30 nations each year. We are extremely grateful
for those who contributed chapters, as well as those who freely gave of their precious
time as reviewers. This book is dedicated to the collegial spirit of scholarship and
camaraderie embodied in the Annual Sea Turtle Symposium.
The Editors
Peter L. Lutz, Ph.D., holds the McGinty Eminent Scholar Chair in Marine Biology
at Florida Atlantic University. Dr. Lutz received both his B.Sc. and Ph.D. from
Glasgow University, Scotland. After finishing his Ph.D. in 1970 he became a research
associate with Dr. Knut Schmidt-Nielsen at Duke University, with whom he worked
on avian respiration. In 1972 he became a Lecturer at the University of Bath,
England, and in 1974 an Associate Professor in Marine Biology at the Rosenstile
School of Marine and Atmospheric Science, University of Miami. In 1982 he was
promoted to Professor and in 1983 became Chairman of Marine Biology and Fish
eries, a post he held until he took up his present position in 1991.
As a comparative physiologist, he has worked on the physiology of a wide
variety of organisms, from liver flukes to duck-billed platypuses. His current interests
center on survival strategies of anoxia-tolerant brains and on the physiology of sea
turtles. Dr. Lutz is a member of the Society for Neuroscience, the American Phys
iological Society, and the Society for Experimental Biology. He is Series Editor for
the Marine Science Series published by CRC Press. He was a Governing Council
Member, Bahamas National Trust, and is a Fellow of the Explorers Club and member
of the Species Survival Commission of the lUCN. He has authored more than 150
research papers and 2 books.
John A. (Jack) Mustek, Ph.D., received his B.A. in Biology from Rutgers University
in 1962 and his M.A. and Ph.D. from Harvard University in 1964 and 1969, respec
tively. He has been on the faculty at the Virginia Institute of Marine Science, College
of William and Mary since 1967 and has successfully mentored 28 M.A. and 30
Ph.D. students. His research has covered many aspects of vertebrate ecology includ
ing community structure in deep sea fishes, shark population dynamics, and sea
turtle ecology. In 1985, he was elected a Fellow by the American Association for
the Advancement of Science. He has served as President of both the American
Elasmobranch Society and the Annual Sea Turtle Symposium and has also served
on numerous national and international advisory groups for conservation and man
agement, most recently on the International Union for the Conservation of Nature
working groups for sharks and sea turtles. He has authored or contributed to more
than 100 research papers and 3 books.
Contributors
Ralph A. Ackerman, Ph.D. Catherine M.F. Lohmann, Ph.D.
Department of Zoolology and Genetics Department of Biology
Iowa State University University of North Carolina
Ames, Iowa Chapel Hill, North Carolina
Karen A. Bjorndal, Ph.D. Kenneth J. Lohmann, Ph.D.

Archie Carr Center for Sea Turtle Department of Biology
Research University of North Carolina
and Chapel Hill, North Carolina
Department of Zoology
University of Florida Molly E. Lutcavage, Ph.D.
Gainesville, Florida Edgerton Research Center
New England Aquarium
Brian W. Bowen, Ph.D. Boston, Massachusetts
Archie Carr Center for Sea Turtle
Research Peter L. Lutz, Ph.D.
and Department of Biological Sciences
Department of Zoology Florida Atlantic University
University of Florida Boca Raton, Florida
Gainesville, Florida
Jeffrey D. Miller, Ph.D.
M.Y. Chaloupka, Ph.D. Queensland Department of Environment
Queensland Department of Environment Townsville, Queensland, Australia
and Heritage
Townsville, Queensland, Australia John A. Musick, Ph.D.
School of Marine Science
Robert H. George, D.V.M. Virginia Institute of Marine Science
Aquatic Animal Medicine Consultants Gloucester Point, Virginia
Gloucester, Virginia
Michael P. O’Connor, Ph.D.
Stephen A. Karl, Ph.D. Department of Bioscience and
Biology Department Biotechnology
University of South Florida Drexel University
Tampa, Florida Philadelphia, Pennsylvania
Colin J. Limpus, Ph.D. David W. Owens, Ph.D.

Queensland Department of Environment Department of Biology
and Heritage Texas A & M University
Brisbane, Australia College Station, Texas
Frank V. Paladino, Ph.D, Michael Salmon, Ph.D.
Department of Biology Department of Biological Sciences
Purdue University Florida Atlantic University
Fort Wayne, Indiana Boca Raton, Florida
Pamela T. Plotkin, Ph.D. James R. Spotila, Ph.D.

Department of Bioscience and Department of Bioscience and
Biotechnology Biotechnology
Drexel University Drexel University
Philadelphia, Pennsylvania Philadelphia, Pennsylvania
Blair E. Witherington, Ph.D.

Peter C.H. Pritchard, Ph.D.
Florida Marine Research Institute
Florida Audubon Society
Department of Environmental Protection
Winter Park, Florida
Tequesta, Florida
Jeanette Wyneken, Ph.D.

Department of Biological Sciences
Florida Atlantic University
Boca Raton. Florida
Table of Contents
Chapter 1
Evolution, Phylogeny, and Current Status............................................................. 1
Peter C.H. Pritchard
Chapter 2
Population Genetics, Phylogeography, and Molecular Evolution......................... 29
Brian W. Bowen and Stephen A. Karl
Chapter 3
Reproduction in Sea Turtles................................................................................... 51
Jejfrey D. Miller
Chapter 4
The Nest Environment and the Embryonic Development of Sea Turtles............. 83
Ralph A. Ackerman
Chapter 5
Orientation, Navigation, and Natal Beach Homing in Sea Turtles..................... 107
Kenneth J. Lohmann, Blair E. Witherington, Catherine M.F. Lohmann,
and Michael Salmon
Chapter 6
Habitat Utilization and Migration in Juvenile Sea Turtles.................................. 137
John A. Musick and Colin J. Limpus
Chapter 7
Sea Turtle Locomotion: Mechanics, Behavior, and Energetics........................... 165
Jeanette Wyneken
Chapter 8
Foraging Ecology and Nutrition of Sea Turtles................................................... 199
Karen A. Bjorndal
Chapter 9
Age, Growth, and Population Dynamics............................................................. 233
M.Y. Chaloupka and John A. Musick
Chapter 10
Diving Physiology................................................................................................. 277
Molly E. Lutcavage and Peter L. Lutz
Chapter 11
Thermal Biology.....................................................................................................297
James R. Spotila, Michael P. O'Connor, and Frank V Paladino
Chapter 12
Hormones in the Life History of Sea Turtles...................................................... 315
David W. Owens
Chapter 13
Salt, Water, and pH Balance in Sea Turtles......................................................... 343
Peter L, Lutz
Chapter 14
Health Problems and Diseases of SeaTurtles...................................................... 363
Robert H. George
Chapter 15
Human Impacts on Sea Turtle Survival................................................................ 387
Molly E. Lutcavage, Pamela Plotkin, Blair Witherington,
and Peter L. Lutz
Index.......................................................................................................................411
1 Evolution, Phylogeny, and
Current Status
Peter C. H. Pritchard
CO NTENTS
1.1 Introduction....................................................................................... 2
1.2 Marine Turtles................................................................................................... 3
1.3 Historical Classifications of Marine Turtles.....................................................5
1.4 Cladistic Placement of Marine Turtle Groups.................................................7
1.5 Marine Turtle Families..................................................................................... 7
1.6 Familial Definitions.......................................................................................... 8
1.6.1 Cheloniidae........................................................................................... 9
1.6.2 Dermochelyidae.................................................................................... 9
1.6.3 Toxochelyidae............................................................................... 9
1.6.4 Protostegidae........................................................................................ 10
1.7 Dermochelyid Diversity................................................................................... 10
1.8 Cheloniid Diversity...........................................................................................11
1.9 Generic Definitions of Living Cheloniids....................................................... 11
1.9.1 Cheionia...............................................................................................12
1.9.2 Eretmochelys....................................................................................... 12
1.9.3 Lepidochelys........................................................................................ 12
1.9.4 Caretta................................................................................................. 12
1.9.5 Natator.................................................................................................13
1.10 Affinities of Recent Cheloniid Species: Current Questions......................... 13
1.11 Survival Status of Living Cheloniids............................................................ 16
1.11.1 Legal and Formal Status..................................................................... 16
1.11.2 Biological Status................................................................................. 16
1.11.2.1 The Green Turtle, Cheionia mydas..................................... 17
1.11.2.2 The Black Turtle, Cheionia agassizii.................................. 18
1.11.2.3 The Flatback, Natator depressus......................................... 18
1.11.2.4 The Loggerhead, Caretta caretta........................................ 19
1.11.2.5 The Hawksbill, Eretmochelys imbricata.............................20
1.11.2.6 The Olive Ridley, Lepidochelys olivácea............................22
1.11.2.7 The Kemp’s Ridley, Lepidochelys kempi............................22
1.11.2.8 The Leatherback, Dermochelys coriacea............................23
References............................................ ...................................................................24
0-8493-8422-2/97/$0.00+$.50
© 1997 by CRC Press, Inc
The Biology of Sea Turtles
1.1 INTRODUCTION
Turtles, being reptiles and tied to the terrestrial environment for oviposition (with
the single partial exception of the chelid Chelodina nigosa [Kennett, 19931), are
generally assumed to have had terrestrial origins. This assumption is probably
correct. Additional evidence is provided by the earliest fossil turtles of the genus
Proganochelys and its relatives that had already achieved a broad distribution in the
northern continents (then still united as “Laurasia”) by the Triassic, and that appear
from their limb structure to have been terrestrial, or, at most, marshdiving forms
(Gaffney, 1990).
Authors such as Römer (1945) and Swinton (1958) assumed that the group of
early reptiles long known as the Cotylosauria, characterized inter alia by the absence
of true temporal fossae, was ancestral to the turtles. Yet intermediate stages were
unknown in the fossil record, the alleged “missing link,” Eunotosaurus Seeley 1892,
from the Permian of South Africa having now been reinterpreted, and Swinton,
although observing that “the Chelonian skull has many features in common with
that of Diadectid Cotylosaurs,” cautiously concluded that “a direct relationship is
not likely.”
Gaffney (1975) considered the turtles to be the sister group of the Synapsida
plus the Diapsida. But he later revised this opinion (Gaffney and Meylan, 1988),
and argued that the turtles were the sister group to the Diapsida alone. Subsequently,
Lee (1993) conducted a cladistic analysis of the Chelonii and various groups of
primitive amniotes and concluded, on the basis of 16 shared derived features, that
the pareiasaurs, a group of anapsid, large, terrestrial reptiles with short, heavy bodies,
and with large osteoderms possibly anticipatory of the chelonian shell, were evolu-
tionarily close to the earliest turtles. The pareiasaurs were, for a time, cosmopolitan
in distribution, but although they flourished in the Late Permian, they were not
persistent. Lee considered the evidence linking the chelonians to other traditionally
cotylosaurian groups such as the captorhinids and the procolophonoids to be weak.
In view of the terrestrial adaptations of the earliest turtles, it is noteworthy that
the subsequent evolution of the Order Chelonii has seen a remarkable proliferation
of aquatic taxa, whereas terrestriality has been retained (or reachieved?), amongst
modern forms primarily by the single family Testudinidae (with about 40 living
species). Some isolated fossil genera (e.g., Meiolania, Zangerlia — see Gaffney
[1983] and and Mlynarski [1972] for discussion) also appeared to have been terres
trial, as are a few living emydids and batagurids. Yet eleven of the twelve living
families are basically aquatic, although in some cases families that are considered
exclusively aquatic today, such as the Pelomedusidae and the Dermatemyidae, once
had significant terrestrial representation (Wood, 1985; Mlynarski, 1972). The great
majority of the extinct families were entirely aquatic also, as far as is known.
Moreover, even specialized aquatic forms, such as the Trionychidae, appeared very
early in the fossil record (Hutchison, 1982). Furthermore, as far back as the early
Jurassic, a newly described turtle, Kayentachelys, from the Kayenta fonnation of
eastern Arizona, considered to be the sister taxon for all other cryptodires, had a
carapace with all of the morphological features of modern, fully aquatic species
(Gaffney et al., 1987).
Evolution, Phylogeny, and Current Status 3
The successful penetration of aquatic niches by both early and modern turtles
was probably made possible by a remarkable example of preadaptation. Other living
aquatic reptiles, including the sea snakes and other snake species adapted to various
degrees for freshwater environments, marine iguanas, crocodilians, etc., as well as
the extinct ichthyosaurs, swim (or swam) by means of body and tail undulations not
dissimilar to those of typical fishes. Turtles, on the other hand, lost the capacity for
this form of propulsion when they developed the shortened, rigid body form and
corselet that has characterized the group since the Triassic. This body form offered
armored resistance to attack by predators, but the tradeoff was reduced speed and
agility, obliging those terrestrial chelonian species surviving in a world with increas
ingly sophisticated predators to adopt specialized (fossorial, cryptic, insular) life
styles (Pritchard, 1979).
Nonetheless, while the encarapaced, terrestrial body form with columnar, walk
ing-type limbs precluded aquatic locomotion by serpentine undulation, it was
remarkably preadapted for very different modes of propulsion, namely swimming
by means of alternating thrusts of limbs with webbed digits, or simultaneous strokes
with powerful paddlelike forelimbs, and these occurred repeatedly.
The shell required only modest modifications in the transition to aquatic life,
these generally taking the form of an overall lower, more streamlined profile, with
sharp, tapered edges to the carapace margins, and expansion of the plastral lobes,
thus facilitating lateral (swimming) rather than ventral (walking) limb movements
(Gaffney et al., 1987).
Only the limbs required profound modifications, and these were indeed forth
coming, the precise form of the newly reshaped limbs being governed by the degree
of aquatic specialization appropriate to the new life-style. In the vast majority of
aquatic chelonians, the limbs represent a compromise between the needs of swim
ming and of walking. In most swimming fonns the hind limbs are longer, more
powerful, and have more extensive webbing than the forelimbs, and they generate
the principal propulsive force, although their narrow specialization for this function
alone is constrained by the need in all chelonian species to utilize the hind limbs
for nest construction.
But in some of the most aquatic chelonians, the forelimbs provide the principal
propulsive force. A trend in this direction is evident in the soft-shelled turtles
(Trionychidae), in which the forelimbs have become substantially modified into
swimming paddles, and the foreclaws (and hindclaws) are reduced to three on each
limb. In these turtles, propulsive force is generated to a comparable degree by
forelimbs and hindlimbs. Forelimb specialization for swimming is carried further in
the family Carettochelyidae (now monotypic, but with an elaborate fossil record and
forebears widespread in northern continents [Mlynarski, 1976]), in which the fore
limbs are paddlelike and two-clawed, but still flexible.
1.2 MARINE TURTLES

Modification of the forelimbs for swimming reached its apogee in the modern marine
turtles of the families Cheloniidae and Dermochelyidae, in which the paddlelike
forelimbs achieve a substantial degree of rigidity by having elongate phalanges
enmeshed in a continuous matrix of tough connective tissue. This not only makes
independent mobility of individual digits impossible, but flexion of the blade of the
paddle as a whole is tightly constrained by the combination of the fibrous binding
of the digits and nonalignment of the corresponding joints between the individual
phalanges of adjacent digits (Zangerl, 1980). In such forms, the forelimb claws are
essentially rudimentary, reduced to one, or two at most. But the claw on the first
digit has a specialized secondary function in adult male cheloniids, in which it is
enlarged and hooklike, and is used for clasping the anterolateral marginal area of
the female during copulation.
Such limbs are poorly adapted for terrestrial locomotion in that the proximal
parts of the limbs bear almost the entire weight of the animal, but ability to locomote
on land cannot be compromised beyond a certain point because of the inescapable
marine chelonian committment to terrestrial oviposition. On the other hand, the
extraordinary demands for superior swimming ability and endurance made by a
marine life incorporating transoceanic migrations ensure that adaptations for aquatic
locomotion are paramount and uncompromised, whereas terrestrial locomotion can
be allowed to become slow and labored, as long as it is not entirely precluded.
Turtles are an important component of marine ecosystems — primarily tropical
and to a lesser degree subtropical ones, but with one species (Dermochelys coriacea)
showing remarkable adaptations for survival and function in very cold water (Frair
et ak, 1972; Greer et al., 1973). Today, however, only seven or eight species of
marine turtles survive, the majority distributed unevenly through all three tropical
oceans, but with three having relatively restricted distributions (the flatback, Natator
depressus, in northern Australia; Kemp’s ridley, Lepidochelys kempi, in the Gulf of
Mexico and North Atlantic; and the black turtle, Chelonia agassizH, in the eastern
Pacific).
Despite this paucity of species, the living marine turtles are not a relictual group.
They have great economic value as well as extreme vulnerability to mankind, at
least while nesting, and their inclusion on most lists of threatened or endangered
species is a reflection primarily of past overexploitation and current need for better
management rather than to inherently poor adaptation to post-Pleistocene conditions.
The documented great diversity of sea turtle taxa in the past probably derives
from several causes, including on the one hand the relatively good chances of
fossilization of these heavy-boned animals, and on the other hand the disappearance
of entire isolated oceans (such as the Niobrara Sea), once populated by diverse and
remarkable sea turtle species. Moreover, the overall evolutionary history of the group
is paralleled on a mega-scale by the history of life itself (Gould, 1980) or on a
human-economy scale by the reduction in the number of automobile manufacturers
in the U.S. during the last century even as cars have become more, rather than less
popular (Yates, 1996). In these examples, an early blossoming forth of numerous
fundamentally “experimental” types (Pritchard and Trebbau [1984] list 27 entirely
extinct genera in the Cheloniidae alone) became winnowed down, probably through
elimination of overspecialized types with poor response to changing conditions, to
a much-reduced diversity of modem survivors, characterized to a considerable degree
by being less rather than more specialized than their forebears (Zangerl, 1980).
Evolution, Phylogeny, and Current Status
1.3 HISTORICAL CLASSIFICATIONS OF MARINE

TURTLES
Linnaeus (1758) included all of the turtles and tortoises that he recognized in the
single genus Testudo, a taxon now restricted to a small group of circum-MediteiTa-
nean terrestrial species of the family Testudinidae. Subsequent early taxonomists in
this pre-Darwinian period failed to identify trenchant carapacial characters to justify
recognition of major divisions of chelonians, and this is understandable in that the
carapace superficially presents extraordinary conservatism. For example, the vast
majority of living and extinct chelonian taxa show the same number of vertebral
and costal scutes (five, and four pairs, respectively), as well as a similar suite of six
pairs of plastral scutes, the commonest variation being in the presence or absence
of an unpaired anterior scute, the intergular.
Turtles and tortoises, as a whole, clearly present a “natural group” (to use the
language of the pre-Darwinians), or a “monophyletic group” or “clade” (to use a
contemporary evolutionary concept), although different opinions have been
expressed on their “affinities” over the last two centuries. (MacLeay, in Jenyns
[1835], perceived or fancied a relationship between the chelonians and the cepha-
lopods, although Jenyns allowed that “the hiatus occurring between is very consid
erable”; and I once saw armadillo specimens stored in the same cabinet as the Bell
collection of chelonians in the attic of the Oxford University Museum, although
there was no one to interpret the implications of this juxtaposition.) Perhaps the
most remarkable “pseudoturtle” that has been described was the Triassic placodont
sauropterygian Placochelys — even its scientific name is turtlelike. This marine
reptile had an oval carapace studded with conical osteoderms, cheloniid-style flip
pers, a head with a narrowed, toothless “beak” (although flattened, shell-crushing
teeth were present on the maxillae and palatines), and reduction of the presacral
vertebrae to about 22 (18 in true turtles). A reconstruction of Placochelys was even
featured, as a sea turtle, in a Hungarian postage stamp in 1969. But the presence of
a single pair of well-developed temporal fenestra high up on each side of the skull,
and many fundamental structural differences in the carapace, make clear that parallel
evolution rather than relationship is the explanation (Römer, 1945, 1956).
The turtles have long been recognized as an order of the class Reptilia, and the
most commonly utilized names for that order include the pre-Linnaean Testudinata
(Klein, 1751), first adopted post-Linnaeus by Oppel (1811); Testudines (Wagler,
1830); and Chelonii (Brongniart, 1800). Bour and Dubois (1985) presented a well-
argued case for utilization of Chelonii as the valid ordinal name today, although this
has not had universal acceptance. Other names that had little or no subsequent use
include Fornicata (Haworth, 1825); Sterrochrotes (Ritgen, 1828); and Tylopoda
(Meyer, 1849).
The first multigeneric system of classification proposed for the Testudines was
that of Brongniart (1805), who considered habitat to be the key character, and
recognized Testudo for the terrestrial species, Emys for the freshwater ones, and
Chelonia for the marine ones. Oppel (1811) presented a dichotomous division of
the turtles at the subordinal level, reeognizing the Chelonii with oarlike forelimbs
and the Amydae with distinct digits. The same division with different names was
used by Merrem (1820) and Bell (1828), who called the marine species with oarlike
limbs Pinnata, and the species with distinct digits Digitata.
Further refinements followed. It was soon recognized that the limbs of tortoises
were as distinct from those of freshwater species as were the limbs of the marine
forms. Moreover, the leatherback turtle and also the freshwater softshells presented
radical distinctions from the typical chelonian body form. This led to Gray’s (1825)
recognition of five families, the Cheloniadae, the Sphargidae, the Trionicidae, the
Emydidae, and the Testudinidae.
Agassiz (1857) reviewed subsequent early nineteenth century classifications,
remarking sagely that the schema of Brongniart, Ritgen, Wagler, Duméril and
Bibron, and Prince Canino Bonaparte, although utilizing totally different names, had
a great deal in common. It was Wagler (1830), followed by Duméril and Bibron
(1836) who hit upon the character that is considered to be of the most fundamental
importance today, namely the plane of retraction of the neck. This feature not only
reflects in the articulations of the cervical vertebrae, but also correlates with impor
tant skull features (such as the trochlear mechanism for the jaw retractor muscles),
and shell characters (the pelvis fused to both carapace and plastron in the pleurodires,
and the presence of an intergular scute in the pleurodires, but only rarely in the
cryptodires). Many pleurodire groups also show remarkable parallelisms with cryp
todires, for example in the loss of the mesoplastra, as well as in the characteristic
carapacial scute mosaic.
Agassiz also commented that many of the classifications that he had reviewed
assumed (implicitly or explicitly) that the turtle species with oar- or paddlelike limbs
were “inferior” to those with separate digits, which in turn were “inferior” to those
with clublike walking feet. This evaluation of his predecessors’ beliefs was correct,
and indeed the assumption contains some elements of truth if we substitute the word
“primitive” for “inferior.” For example, the marine turtles do show the complete
(nonemarginate) skull roof considered characteristic of early turtles, and the char
acteristic elephantine foot of the testudinids is indeed probably a derived rather than
a primitive feature. But it should not be forgotten that both Agassiz and the ante
cedents whom he quoted were nonevolutionists, and thus concepts of primitive vs.
advanced features, or of “natural groups,” had at most a theological basis rather than
one based upon actual closeness of phylogenetic relationship.
Subsequent classifications, of living turtles at least, were based upon the primary
division of turtles and tortoises into Pleurodires and Cryptodires, although the proper
alignment of the leatherback and the softshells remained a challenge. The many
morphological peculiarities of the former (summarized by Gervais, 1872) led Cope
(1872) to declare this taxon the “sister group” (to use cladistic terminology) for all
other turtles. He named it “Athecoidea” (or, in family-level nomenclature, “Athe
cae”), meaning “without a shell.” This allocation was followed by Dollo (1886),
Boulenger (1889), Lydekker (1889), and various others, the latest author of note to
use it being Carr (1952). In these classifications, all other chelonians were identified
as Thecophora (“shell-bearing ones”).
Furthermore, the Thecophora were trichotomously divided into the superfamilies
Cryptodira, Pleurodira, and Trionychoidea (Boulenger, 1889). Such elevation of
status of the taxa with scuteless shells is understandable, in that it makes a mockery
of classifications founded upon details of scutation when some turtles have no scutes
at all. But, despite their extremely divergent morphology, Dermochelys and its fossil
relatives are now considered to be related to the marine turtles of the family Che-
loniidae (Gaffney and Meylan, 1988).
Yet Cope’s early (1872) designation of a taxon (Athecoidea) linking the pro-
tostegids and the dermochelyids, while shaken by Zangerl’s subsequent clarification
that protostegids (in contrast to demiochelyids) had normal chelonian scutes rather
than leathery skin, achieved some vindication with Gaffney and Meylan’s (1988)
proposal of an “Epifamily” Dermochelyoidae, with the same content. Moreover,
even Dollo, while recognizing the Athecae on the basis of its extreme morphological
modifications, considered it to be derived from the Cheloniidae (Dollo, 1886).
Dollo’s conclusion may be factually correct, although the divergence was so far back
that it is arguable whether or not the common ancestral stock was truly cheloniid.
But his nomenclatural inteipretation of its implications, although permissible by
classical taxonomists, is in conflict with the rules of Cladism, which seeks to establish
a nomenclature based purely upon temporal order of divergence rather than upon
degree or extent of divergence.
Today the trionychids also are recognized only as a family, related to the caret-
tochelyids, and incorporated within the Cryptodira (Meylan, 1987).
Various earlier, intermediate, or conflicting placements of the Dermochelyidae
are discussed by Pritchard and Trebbau (1984).
1.4 CLADISTIC PLACEMENT OF MARINE TURTLE

GROUPS
Gaffney and Meylan (1988) offer the following cladogram (Figure 1.1) for the marine
turtles with well-developed flippers (i.e., excluding the Thalassemyidae).
While I have many misgivings about the cladistic process as a whole (Pritchard,
1994), this cladogram has merit in that it recognizes areas of uncertainty without
attempting arbitrary resolution, including the familial placement of Notochelone,
Allopleuron, and Erquelinnesia, and the unresolved trichotomies for the osteopygid
genera as well as the Notochelone complex. It is also clearly preliminary in that, of
the 31 cheloniid or possibly cheloniid genera mentioned by Pritchard and Trebbau
(1984), only nine are mentioned. There have also been recent discoveries clarifying
the relationships between the extant cheloniid species (Bowen et al., 1993) that are
at variance with the cladogram.
1.5 MARINE TURTLE FAMILIES

Marine adaptation has been undertaken more than once in the history of the Chelonii.
The earliest marine turtles appeared in the Jurassic, with a marine species of the
family Pleurosternidae, Desmemys bertelsmanni, and an entirely marine family, the
Thalassemyidae also appearing in the Jurassic, possibly derived from the primitive,
probably freshwater Plesiochelyidae. The Thalassemyidae showed many carapacial
FIGURE 1.1 Cladogram “D”, Chelonidea.
parallels (or anticipations) with later marine turtle families, but the limbs remained
unspecialized for marine life (Hay, 1908), and Gaffney and Meylan (1988) recently
found little or nothing to separate Thalassemys itself from the plesiochelyids. Cer
tainly, Hay’s concept of the Thalassemyidae was a polyphyletic one, with his family
definition following rather than preceding the allocation of a mixed bag of genera
from at least three distinct families.
Some extinct Pelomedusid genera (e.g., Taphrosphys [Wood, 1974]; possibly
Stupendemys [Wood, 1976]) may also have been marine.
By the Cretaceous, four marine turtle families — the Toxochelyidae, Protoste-
gidae, Cheloniidae, and Dermochelyidae —were all established, the last two surviv
ing to the present. Whether or not these all represent independent invasions of the
marine environment is unclear, but it is probable (as Dollo observed) that the
dermochelyids represent an extraordinarily divergent offshoot of the cheloniids,
whereas some of the toxochelyids bear such a strong superficial resemblance to
certain cheloniids that the possibility of close relationship must be seriously enter
tained.
1.6 FAMILIAL DEFINITIONS

Detailed definitions are possible for the living marine turtle families (Pritchard and
Trebbau, 1984), although the proviso has to be stated that the applicability of soft-
tissue characters, established from the living taxa, to the fossil representatives is
obviously conjectural. Gaffney and Meylan (1988) give diagnoses for these families
based upon purely osteological features. Definitions for the entirely extinct marine
families are necessarily less complete.
1.6.1 C heloniidae
A family of turtles characterized by an extensively roofed skull with well-developed

rhamphothecae; secondary palate present; incompletely retractile or nonretractile
head; extremities in the form of nonretractile flippers covered with numerous small
scales; the forelimbs having highly elongate digits firmly bound together by con
nective tissue, the claws being reduced to one or two on each limb, and the radius
and ulna immobilized against independent movement by juxtaposed rugose surfaces;
shell covered with horny scutes, variable in number, but commonly including five
vertebráis and six pairs of plastral scutes, together with an unbroken series of three
or four pairs of inframarginals, multiple axillary scutes, and usually both an interguiar
and an interanal scute; the plastron often with persistent fontanelles, one in the
middle and others in the entoplastral and xiphiplastral regions; hyoplastra and hypo-
plastra not suturally connected mesially, but each with a series of coarse spikes that
interdigitate mesially; plastron not crucifonn, and posterior plastral lobe relatively
long and wide.
1.6.2 D ermochelyidae
A family of turtles characterized by extreme reduction of the bones of the carapace
and plastron (with the neural and peripheral bones of the carapace, and entoplastron
of the plastron, lacking; the pleurais reduced to endochondral ribs, separated by
wide fenestrae; and the plastral bones reduced to narrow splints, forming a ring of
bones surrounding a great fontanelle); development of a neomorphic epithecal shell
layer consisting of a mosaic of thousands of small polygonal bones; claws and shell
scutes lacking (scales only present in the first few weeks of life); skull without nasal
bones; jaw surfaces covered with keratin, but lacking differentiated rhamphothecae;
parasphenoid overlain by pterygoids; prefrontals in contact dorsally, with descending
processes that are moderately separated; unridged tomial surfaces; a generally neo-
tenic and oil-saturated skeleton; extensive areas of vascularized cartilage in the
vertebrae, limb girdles, and limb bones; very large body size; and marine habitat.
1.6.3 T oxochelyidae
The toxochelyids were a group of small to medium-sized, broad-shelled (circular
to cordiform) marine turtles that diverged from the early cheloniid stock in the early
Cretaceous and persisted in the proto-Atlantic and Niobrara seas until the late
Eocene. Various toxochelyid genera display progressive development of a secondary
palate, from Toxochelys with a primary palate only, through Ctenochelys with
incipient undershelving, a more advanced condition in Osteopygis, and extreme
development of the secondary palate in Erquelinnesia. The carapace ranged from
fully closed (i.e., without intercostal fontenalles) in Osteopygis to a highly reduced
condition with persistent, large fontanelles in Erquelinnesia. Features characteristic
of the family include the presence of small nasal bones and the well-developed
10 The Biology of Sea Turtles
peripherals (often reduced to narrow bars in cheloniids). The plastron, primitively

cruciform, was larger and with much wider bridges in later representatives. In one
subfamily (the Lophochelyinae), the neural ridge was highly developed and serrated
or tuberculate in lateral aspect. The toxochelyids differed from the thalassemyids
not only in the cruciform plastron, but also in having narrow vertebral scutes (much
narrower than the pleurais); unreduced neurals; and two suprapygals. In many
features of the skull they appeared intermediate between cheloniids and chelydrids
(i.e., snapping turtles). The toxochelyids are discussed by Hay (1908) and Zangerl
(1953, 1980).
1.6.4 Protostegidae
The protostegids were a group of large to gigantic turtles whose more primitive
members (e.g., Rhinochelys) have been found in Cretaceous deposits of France and
England, but which reached its maximal development (with the genera Protostega
and Archelon, known from Upper Cretaceous deposits of Texas, South Dakota,
Arkansas, Alabama, Colorado, and Kansas) in the Niobrara Sea. They showed certain
parallels with the dermochelyids, one of these being the possession of vascularized
chondroepiphyses (Rhodin, 1985), a feature present in a number of fossil turtle
genera (Psephophorus, Eosphargis, Archelon, Pneumatoarthrus, Corsochelys) as
well as the living Dermochelys. However, vascular chondroepiphyses, a condition
readily evident in the ends of the limb bones even of purely fossil material, may be
a parallelism connected with very large adult size (Pneumatoarthrus was originally
described as a dinosaur! [Cope, 1870; Baird, 1978]), and rapid growth potential
rather than an indicator of relationship.
The protostegids were characterized by large skulls (up to 100 cm in length in
the case of Arche Ion) with well-developed, sometimes strongly hooked beaks; no
secondary palate; reduced xiphiplastra and reduced or absent epiplastra; considerable
reduction of the carapacial armor, in some cases with intercostal fontanelles reaching
close to the vertebral column even in adults; a tendency towards keeled or tuberculate
middorsal keels (sometimes with carination present only on alternating neural
bones); and a slight to moderate degree of temporal emargination.
1.7 DERMOCHELYID DIVERSITY

The single extant dermochelyid, D. coriacea, shows very modest geographic vari
ation, and it is probable that no subspecies exist (but see Brongersma [in press] for
a discussion of potentially available names for different populations). Dermochelyids
appear not to fossilize well or easily, and the majority of the rather few fossil genera
reviewed by Pritchard and Trebbau (1984) are known from fragmentary material
only. For example, Cosmochelys (Eocene of Nigeria) is based upon fragments of
the epithecal carapace, and is not even considered dermochelyid by some authorities,
although it is so accepted by de Broin and Pironon (1980). Pseudosphargis (Late
Oligocène of Germany) is known from a cranial fragment only, and most species
of the polytypic Psephophorus (Eocene-Pliocene, Europe, North Africa, North
America) are known from shell fragments only, although P. rupeliensis is known
from several mostly complete specimens. In this genus, the shell is characterized by
thickened, strongly keeled epithecal bones that form a complete layer on the plastron
as well as the carapace (in contrast to Dermochelys).
In the genus Eosphargis (Eocene, Europe), a well-preserved skull shows typi
cally dermochelyid foim, but the mosaic epithecal layer was not yet developed
(Nielsen, 1959).
1.8 CHELONIID DIVERSITY

Pritchard and Trebbau (1984) inventoried 31 cheloniid genera (only four with living
representatives — Natator was resurrected later). However, many of these were
known from fragmentary material only — some only from a mandible, or a humerus
— and, despite the abundance of named forms, we remain a great distance from a
full comprehension of the phylogeny and morphological diversity of the family
Cheloniidae.
Probably the best brief summary is that of Zangerl (1980). He characterized
cheloniid evolution as a persistent theme of specialized forms that evolved, but failed
to persist, whilst a generalized stem group, adapted for near-shore rather than pelagic
waters, survived and progressively generated subsequent short-lived, relatively
pelagic, specialized offshoots. Thus, in the Cretaceous, named offshoots included,
on the one hand, Glyptochelone suyckerbuyki, and on the other hand a progression
of forms (Catapleura arkansaw; Desmatochelys lowi; Corsochelys haliniches; and
Allopleuwn hoffmarmi) that showed increasing pelagic specialization. Zangerl con
sidered pelagic adaptation to be manifest in elongate, persistently fontanelled cara
paces (as opposed to broad, fully ossified ones), and the plastron also with persistent
median and lateral fontanelles. Other specializations include the development of a
secondary palate and the conversion of the hind limbs into relatively stiff “rudders,”
rather than active propulsive members. But the situation is complex; many forms
show a mosaic of primitive and advanced characters, and one extant species (L.
oliváceo) has a fundamentally primitive shell, with broad, short outline and virtually
complete ossification, yet is so pelagic that the juveniles are almost never found,
despite this being the most abundant surviving marine turtle species.
Eocene cheloniids include Puppigerus camperi; Argillochelys subscriptata; and
Eochelone brabantica, only the last being considered to show pelagic specialization.
In the Oligocène, named taxa include Procolpochelys sp.; Oligochelone rupeliensis;
and ''Chelonia’' gwinneri. Miocene forms include P. grandaeva, P. melii, and Syl-
lomiis aegyptiacus. The only well-documented Pliocene species is Chelonia sismon-
dai.
1.9 GENERIC DEFINITIONS OF LIVING CHELONIIDS

It is easier to write a generic diagnosis for a large, polytypic genus than for a
monotypic one. For the former, a list of common features of the various recognized
species will suffice. However, for the latter, it is necessary to separate those char
acteristics that merely characterize the single known species from “true” generic
characters, that would permit correct diagnosis and allocation of a new potential
congener, fossil or living, that was unknown when the diagnosis was drawn up.
The diagnoses of Carr (1942) for four of the known cheloniid genera meet these
criteria, and are quoted below with very minor modifications. On the other hand,
the two recently presented and much more detailed diagnoses for Natator (Zangerl
et al., 1988; Limpus et al., 1988) are indistinguishable from diagnoses of the single
species N. depressus, in that they combine both fundamental (generic-level) and
species-specific data.
1.9.1 C helonia
One pair of prefrontal scales; homy cutting edge of lower jaw coarsely dentate, that
of upper Jaw strongly ribbed vertically; bony alveolar surface of upper jaw with a
low, but regularly raised auxiliary ridge behind the anterior ridge which is very
strong and terminates anteriorly in a pointed eminence at the posterolateral corner
of the premaxillary pit; costal scutes 4 pairs, well cornified, juxtaposed.
1.9.2 Eretmochelys
Two pairs of prefrontal scales; nuchal scute not in contact with first costáis; snout
elongate, narrow, the mandibular symphysis deeply excavated and not terminally
toothed; bony alveolar surface of upper jaw with a sharp-crested ridge; dorsal scutes
usually 4 pairs, usually thick and conspicuously imbricated; peripheral bones usually
11 (after Carr, 1942).
1.9.3 L epidochelys
Maxillaries not in contact, separated by vomer; frontal bone usually entering rim of
orbit; pterygoids markedly broadened anteriorly, the ectopterygoid processes strong;
fontanelles in choanal chamber near opening, not hidden by alveolar surface in
ventral aspect; external openings of orbits not concealed by overlying bones in
ventral aspect; descending processes of prefrontals not reaching palatines; lower jaw
with a more or less sharp and strong median elevation at the posterior border of the
bony alveolar surface, which may or may not extend forward as an elevated ridge;
four enlarged inframarginal scutes on the bridge; costal scutes thin, juxtaposed, 5
to 9 pairs (often asymmetrical); neural bones 11 to 15.
1.9.4 C aretta
Maxillaries in contact, not separated by vomer; pterygoids not, or but slightly,
broadened anteriorly, the ectopterygoid processes vestigial or lacking; fontanelles
far forward in choanal chamber, completely concealed by alveolar surface in ventral
aspect; external openings of orbits concealed by overlying bones in ventral aspect
(in mature specimens); descending processes of prefrontals connected with palatines;
bony alveolar surface of lower jaw smooth at posteromedian border; symphysis of
mandibles without longitudinal ridge; bridge with three enlarged inframarginal
scutes; costal scutes well comified, juxtaposed, usually five pairs; neural bones
usually 7 or 8.
1.9.5 N atator
Maxillaries widely separated by vomer; pterygoids broadened anteriorly and with
very strong ectopterygoid processes; fontanelles far forward in choanal chamber,
completely concealed by alveolar surface in ventral aspect; external openings of
orbits not concealed by overlying bones in ventral aspect; symphysis of mandible
with a strong median ridge terminating posteriorly in an elevated prominence;
external openings of internal carotid canals very large; head scalation characterized
by a pair of preoculars lying between maxillary rhamphotheca and prefrontals, post-
parietals variable and often asymmetrical, generally including an undivided median
element; plastral fontanelles closing at sides, but remaining open medially at matu
rity; four pairs of inframarginal scutes; carapacial scutes very thin, poorly kerati
nized, and with sulci tending to disappear with age; costáis four pairs. Neural bones
about twelve, with frequent transverse division of “standard” elements, and 2 or 3
suprapygals.
From the above diagnoses, the confusing “mosaic” of characters of Natator is
evident. The skull is so similar to that of L. kempi that some of the older skulls in
Australian museums have been identified and catalogued as that species.
1.10 AFFINITIES OF RECENT CHELONIID SPECIES:

CURRENT QUESTIONS
With only six or seven extant cheloniid species, the relationships between these
species have been surprisingly conjectural until very recent times. The key questions
may be phrased as follows:
1. What subfamilies (or “Tribes”) should be recognized, and what is the

content of each of the recognized subgroups?
2. What is the correct placement of the flatback turtle?
3. What is the relationship between the two forms of ridleys?
4. What is the status of the “black turtle” of the eastern Pacific?
Deraniyagala (1939) recognized a superfamily, the Chelonioidae, comprising

two families, the Cheloniidae (Chelonia and Eretmochelys), and the Carettidae
{Carena and Lepidochelys). The latter was distinguished by the greater number of
certain shell elements, specifically the peripheral bones and the costal scutes.
(Deraniyagala considered that four pairs of costáis and eleven pairs of peripheral
bones constituted a phylogenetieally reduced count, whereas today the proliferation
of elements beyond these numbers is considered to be the derived condition.)
Carr (1942) disagreed shaiply with Deraniyagala, in a paper that uneannily
anticipated the cladistic process. Using 25 characters (some not independent, and
unanalyzed in terms of primitive vs. derived), he found only four that justified placing
Eretmochelys with Chelonia, and 15 justifying placement of Eretmochelys with

Caretta and Lepidochelys. He concluded that the Cheloniidae should include only
Chelonia, the other three genera being placed in the Carettidae. Deraniyagala (1953)
partially agreed, or at least was willing to meet Carr half way, and recognized the
subfamilies Cheloniinae, Carettinae, and Eretmochelinae, all within the family Che
loniidae.
Subsequent authors were divided as to whom to follow. Frair (1979) and Prit
chard and Trebbau (1984) favored Carr (1942); Zangerl and Turnbull (1953), Mly-
narski (1969), and Smith and Smith (1980) followed Deraniyagala (1939), although
Smith and Smith overstated their case by claiming that it was “universally recog
nized.” Nobody followed Deraniyagala’s 1953 classification, although it was not
totally without merit.
Using genetic techniques unavailable to earlier workers, Bowen et al. (1993)
found that all phylogenetic analyses of mtDNA data supported placement of the
hawksbill with the Carettinae, suggesting that the unusual spongivorous diet of the
hawksbill probably evolved from a carnivorous rather than an herbivorous ancestral
condition.
The flatback turtle was originally described as Chelonia depressa by Carman,
1880. The type series was composite (including a specimen of C. mydas), but the
lectotype (MCZ 4473) was recognized by Carr (1942) as “an extraordinary speci
men,” with characters reminiscent of both Chelonia and Lepidochelys. Baur (1890)
thought that depressa belonged in its own genus, and McCullough (1908) agreed,
naming the new genus Natator, in combination with a new specific name, tessellatus.
Fry (1913) also agreed that the flatback was a distinctive species, and published a
detailed description. However, all of these works were generally ignored until Wil
liams et al. (1967) formally resurrected C. depressa as a valid cheloniid species.
Subsequent morphological analyses (Zangerl et al., 1988) and both electro
phoretic and osteological comparisons (Limpus et al., 1988) demonstrated conclu
sively that the flatback should be recognized as a distinct genus, Natator, with closer
affiliations with the Carettini than with the Chelonini. Limpus et al. did not examine
L. kempi or C. agassizi, but they presented a phylogeny of the remaining species,
based upon electrophoretic data, that placed Natator and L. olivácea close together,
Caretta and Eretmochelys in a separate branch and more distantly related, and
Chelonia mydas as the sister group to the other four taxa. However, Bowen et al.
(1993), using mtDNA analysis, found that Natator had comparably wide genetic
separations from both the Chelonini and the Carettini. The only conclusion one can
draw from this is that the flatback is the only extant representative of a new tribe,
the Natatorini.
Incidentally, in that the gender of Natator is not self-evident, it must be construed
as masculine because its describer (McCulloch, 1908) used it in combination with
a masculine-form epithet {‘‘tessellatus"'). The correct name for the flatback turtle is
thus N. depressus.
When examined side by side, the close relationship between the two species of
ridley (L. olivácea and L. kempi) is obvious. Indeed, there is no external difference
between a normal juvenile L. kempi and a specimen of L. olivácea with only five
pairs of costal scutes, a minority condition, but not rare in some areas (Pritchard,
1969). However, earlier workers lacked the material to make these comparisons, and
the two were placed in separate genera {Colpochelys and Lepidochelys) for a long
time. Carr (1942) had to report that prevailing opinion even then insisted either that
the two ridleys and the loggerhead should each be in separate genera, or that all
three forms should be united within Caretta. Yet, he added, it “seems evident that
kempii should be associated with olivácea in the genus Lepidochelys Fitzinger,” an
allocation presaged by Baur (1890) and followed by most subsequent authors (see
Pritchard, 1969, 1989 for details).
But there were holdouts, the most prestigious being Loveridge and Williams
(1957) and Mertens and Wermuth (1955), who considered kempi to be merely a
subspecies of L. olivácea. Once again, Bowen et al. (1993) offered a mitochondrial
DNA analysis, in this case demonstrating that, although the two forms were sister
taxa, they showed genetic distinction at the species level.
The “black turtle” was described as a full species, Chelonia agassizii, by Bocourt
(1868). C. mydas caninegra Caldwell 1962, appears to be a synonym, although the
assumption that this forni from the Gulf of California is identical to agassizii from
Pacific Guatemala has not been proven (but is probable). The pros and cons of
species vs. subspecies status for agassizii have been discussed by Mrosovsky (1983)
and Pritchard (1983). I have justified my own position (full species status) on
narrowly technical grounds (i.e., the nomenclature of the original describer, Bocourt,
should stand until formally changed), and on biological ones, in that agassizii and
mydas-Y\kç^ forms are sympatric in several places in the Pacific, including the Gal
apagos islands and New Guinea; the degree of differentiation in size, shape, and
color is more extreme than that found in any other Chelonia population; the dark
plastral coloration of agassizii is not environmentally derived; and there may be
purely physical reasons why a 40-kg adult male agassizii may be unable to mate
successfully with a 200-kg adult female mydas, even if the two do come in contact.
Subsequent contributions on the subject include that of Kamezaki and Matsui
(1995), who analyzed geographic variation in skull morphology of six Chelonia
populations (two from each tropical ocean), and concluded that the Galapagos
sample (i.e., agassizii) was completely separated from the other samples by a
canonical discriminant analysis, and that this result indicated distinctness of the
eastern Pacific population. On the other hand, Bowen et al. (1993) found that eastern
Pacific turtles represented but a small subset of lineage diversity within the broader
and deeper mtDNA gene tree for the globally distributed green turtle.
A conclusive decision on this issue thus remains elusive. Bowen et al. found
that nDNA data indicated very close relationship between the Galapagos and the
Pacific Mexican populations of "'agassiziC, but both lacked close affinities with
south-central and western Pacific populations, which argues in favor of species-
level status for agassizii. On the other hand, a phylogeny derived from nDNA links
the eastern Pacific populations with, inter alia, those of Ascension Island and the
Atlantic Coast of Africa (Parham and Zug, 1996)! This discordance of nDNA with
both morphology and geographic probability would, to some, substantially lower
the credibility of nDNA analysis as a technique for elucidating phylogenetic
relationships.
Systematists have been divided in recent years both in whether to recognize

subspecies at all, and, if the subspecies concept is to be rejected, whether to “pro
mote” previously recognized subspecies to full species status, or to “demote” them
into oblivion. Widespread, genetically complex species such as Chelonia mydas
present especial challenges when one tries to force them into rigid nomenclatorial
systems, and some of the implications of recognizing agassizii at all (including the
obligation to give nomenclatorial status to many other populations of Chelonia
mydas) are discussed by Parham and Zug (1996).
1.11 SURVIVAL STATUS OF LIVING CHELONIIDS

1.11.1 Legal and Formal S tatus
The U.S. Department of the Interior, under the authority of the Endangered Species
Act, currently lists C. mydas (endangered in Florida and Pacific Mexico) and Caretta
caretta as Threatened Species. E. imhricata, L. olivácea, L. kempi, and Dermochelys
coriácea are listed as endangered. N. depressus, the most restricted and least-known
of all living cheloniids, is unlisted.
The Convention on International Trade in Endangered Species of Flora and
Fauna (CITES) lists all marine turtles on its Appendix I (i.e., prohibited from
international trade from or to signatory countries).
The World Conservation Union (lUCN) has listed C. caretta as Vulnerable and
all other sea turtle species (except A. depressus) as Endangered. However, the lUCN,
in close cooperation with the Secretariat and Parties to CITES, has now adopted a
set of complex numerical and ostensibly objective criteria by which the status
category of a species should be deduced. While these criteria are expected to apply
only to species proposed for the CITES appendices in the future, they will be
incorporated across the board by lUCN. The criteria incorporate considerations of
actual global population numbers, fragmentation of habitat and populations, and
demonstrable population trends. For the great majority of species, the necessary data
are unlikely to be currently available. Among the sea turtles, even the “critically
endangered” L. kempi may only qualify as “conservation dependent,” in view of its
modest population recovery in recent years.
1.11.2 B iologïcal S tatus
The actual, biological status of marine turtles is a topic of considerable complexity.

Unlike, say, endemic species of fauna or flora on small, remote oceanic islands,
where a single ecological perturbation may precipitate a chain of events leading to
multiple extinctions, rather few marine species are faced with biological extinction,
and typically those that have been exterminated, such as Stellers’s Sea Cow or the
Caribbean monk seal, were large, intrinsically vulnerable species with very small
natural ranges.
Rather, marine species are faced with problems of overharvest, or even “popu
lation collapse” and “commercial extinction” at some point short of biological
extinction. In other words, the “demostat” for such species may be drastically shifted
to a lower position once anthropogenic stresses (directed or incidental) become

manifest and unremitting, but, typically, the new setting of the demostat is not at
the zero point. Nonetheless, it remains to be seen whether these conceptual gener
alizations about marine species are applicable to marine turtles, in that the unavoid
able terrestrial ovipositional excursions of all marine turtle species constitute a
special vulnerability to which fully marine species are not subject.
The biological status of the eight (or seven) surviving marine turtle species are
summarized below.
1.11.2.1 The Green Turtle, Chelonia mydas

This is a circumglobal species, most of whose important nesting and feeding grounds
lie within the tropics. It has major nesting colonies on mainland shores (such as
northwestern Costa Rica, or the coast of eastern Surinam), on barrier reef islands
(Queensland, Australia; d’Entrecasteaux Reef, New Caledonia), and on remote oce
anic islands (e.g.. Ascension Island, Atol das Rocas). In many places it has long
been harvested for meat and eggs. Demand for international commerce is now an
insignificant factor, but has been replaced by increasing demand for subsistence and
local markets by indigenous people, whose population increase has often not been
matched by an increase in real wealth or political opportunity.
St",
'■ iT ^ u ‘
FIGURE 1.2 The green turtle, Chelonia mydas.
Thus, it remains to be seen whether the Tortuguero, Costa Rica, nesting colony
of the green turtle, although still large and the most important by far in the Caribbean,
can indefinitely survive heavy subsistence take by Costa Ricans from the city of
Limon as well as by Miskito people from Caribbean Nicaragua, San Bias people
from Panama, and Guajiros from northeastern Colombia and northwestern Venezu
ela. Similar uncertainties are faced by the large Australian colonies, protected whilst
in Australian territory and waters, but heavily harvested in various islands of Indo
nesia (Limpus, 1994, 1995).
Here and there, green turtle colonies seem to be on the increase. In the last
decade, not only has regular green turtle nesting been observed where none was
seen before at Rancho Nuevo, Tamaulipas, Mexico, but also, on the mid-Atlantic
coast of Florida, U.S.A., green turtle nesting is now commonplace on beaches where
almost nothing but loggerheads nested two decades ago. These increases on well-
patrolled beaches seem to be real, and with good protection now offered to many
nesting colonies, including those of Atol das Rocas, Ascension Island, and Trindade,
the green turtle does not appear to be faced with imminent extinction.
Parenthetically, it is interesting to note that proponents of turtle farming and
ranching have observed that, although the green turtle has been sought by interna
tional gourmets for centuries, it is no more endangered than other sea turtle species,
and thus commercial utilization could not have been a significant factor in any
decline that the species may have experienced (Fosdick and Fosdick, 1994).
1.11.2.2 The Black Turtle, Chelonia agassizii
This species (possibly only a subspecies of C. mydas) is confined to the eastern

Pacific. The species is protected in the Galapagos Islands and is nominally protected
in Mexico also, where the important nesting grounds in Maruata Bay are patrolled
by teams from the Universidad de Michoacan. Nonetheless, individuals from both
the mainland and Galapagos nesting grounds are caught in uncontrolled numbers in
Peruvian waters, and are also subject to illegal harvest on the Mexican and Central
American Pacific coasts, including the Gulf of California. Furthermore, marine
conservation efforts in the Galapagos Islands have been subjected to severe chal
lenges by settlers and fishermen in the last two years (Pritchard, 1996).
Thus, although the numbers are probably lacking to establish an unequivocal
status categorization following the new numerical criteria, the outlook for C. agas
sizii is, in the long run, uncertain. Much hinges on the fundamental question of
whether or not total protection of nesting turtles, their eggs, and their nesting beach
can offset uncontrolled harvest on the feeding grounds hundreds or thousands of
kilometers away. Answers to this key question are not yet available.
1.11.2.3 The Flatback, Natator depressus
This species is unlisted by the authorities of USDI and lUCN because it is essentially
confined to the waters of a single nation (Australia), where it is protected by law
except from aboriginal harvest, and where most of the nesting beaches are remote
and undeveloped (with the exception of the easily accessible Mon Repos Beach in
Queensland). Moreover, Australian native people in general prefer to consume the
more abundant and more succulent C. mydas. The outlook for Natator would thus
appear to be good, but the vulnerability of individuals to incidental capture in prawn
trawls, and the overall relatively restricted range lending panspecific vulnerability
from single catastrophic environmental events, leave no grounds for complacency.
FIGUKIC 1.3 "Hie black tuide, Chekmia agassizM.
FIGURE 1.4 The flatback, Natator depressus.
1.11.2.4 The Loggerhead, Caretta caretta

The loggerhead is little sought for its flesh, and although the eggs are gathered in
some parts of the world, direct take for human consumption is not a major factor
in its survival prospects. Rather, this species has an “antitropical” distribution that
not only fragments its overall range into well-separated enclaves in the northern and
southwestern Indian Ocean, eastern Australia, Japan, southeastern U.S., the Medi
terranean, and southern Brazil, but it also brings the species into contact with
industrial and development stresses ranging from massive incidental capture in
Atlantic shrimp trawls to resort and recreational development of nesting beaches.
At present, it appears that nesting populations are declining as a result of incidental
catch in both southern Queensland, Australia (Limpus, 1994) and in the U.S. north
of Cape Canaveral, but on the other hand the larger populations in Florida south of
Cape Canaveral and also the relatively small population in Natal, South Africa, are
increasing (National Research Council, 1990).
FIGURE 1.5 The loggerhead. Caretta caretta.
The small or medium-size populations in the Mediterranean, nesting primarily

in Zakynthos and other islands of Greece and the Turkish coast, and with important
feeding grounds off the coast of Tunisia, are threatened by a variety of factors. These
include intensive touristic development of many of the best nesting beaches. On the
other hand, recent information (Venizelos, 1996) identifies by far the most important
loggerhead nesting beach in the entire Mediterranean in eastern Libya, with an
estimated 9000 nests along 1250 km of coast, and with no foreseeable prospects for
touristic development.
1.11.2.5 The Hawksbill, Eretmochelys imbricata
Intense national and international trade in hawksbill shell products and, in more
recent decades, entire stuffed hawksbill turtles, has led to widespread concerns that
the species is being seriously overexploited. Moreover, although this may be the
most commonly seen species in certain tropical reef habitats in the Caribbean Islands
or tropical Australia, such populations generally consist primarily of subadults, and
very few nesting “colonies” (as opposed to isolated nesting females) are known.
Certain!; it is fundamentally less abundant than the green turtle, but downward
The green turtle
t i* Chelonia m ydas
"'J' '.'T i
The black turtle

Chelonia a g a ssizii
The flatback
Natator depressus.
The loggerhead
Caretta caretta
The hawksbill
Eretm ochelys im bricata
Olive ridley
Lepidochelys oUvacea
Kemp’s ridley
Lepidochelys kern p i
The leatherback
D erm ocheiys coriacea
trends are often difficult to determine from available data, and some of the more
dire warnings about imminent extinction of the species have stemmed simply from
the contrast between the very substantial global harvest for shell and other products,
and the modest size of the known nesting populations. In a global analysis of the
status of the hawksbill, Groombridge and Luxmoore (1989) could only classify it
as “indeterminate.”
FIGURE 1.6 The hawkshill, Eretmochelys imbricata.
For many decades, Japan has constituted the largest importer of raw tortoiseshell.
However, Japan has now withdrawn its reservation to this species on the CITES
convention, where it is listed as Appendix I (banned in international commerce),
and has phased out importation of hawksbill products. On the other hand, Cuba —
a major habitat for the hawksbill and supplier of world tortoiseshell markets — has
recently expressed the opinion that its domestic hawksbills not only constitute a
nonmigratory stock, but also have a sufficiently healthy population to justify down
grading to CITES Appendix II. The intention is to develop hawksbill ranches in
Cuba to supply the Japanese demand. The implications of this move are ambiguous,
but the conditions for approved commercial ranching of Appendix II species agreed
by the Parties to CITES in November 1994 are sufficiently demanding that a legal
Cuban turtle ranching program may well be linked to much better protection of
Cuban, and regional, hawksbill stocks than has hitherto been available.
In a review of worldwide population trends of the hawksbill, Limpus (1995)
could only find one example of an increasing hawksbill population — that of the
Sabah Turtle Islands (Malaysia), where a more than tenfold increase has been logged
since 1969 (Trono, 1994).
1.11.2.6 The Olive Ridley^ Lepidochelys oUvacea

The olive ridley, although having relatively localized nesting, remains the most
numerous species of sea turtle in the world as a result of the continued existence of
a few sites of enormously aggregated nesting — two in Pacific Costa Rica, one in
Pacific Mexico, and two or three in northeastern India, with some more minor sites
in Nicaragua and scattered nesting along certain other tropical mainland shores.
Whether or not the existence of such numbers of turtles on these few nesting beaches
is reason to believe that no problems exist is debatable. In all cases except for the
limited egg harvest program at Playa Ostional, Costa Rica, these “arribada” beaches
are nominally protected, although incidental take by trawlers is significant in both
Costa Rica and India, and the Indian sites may also be threatened by fishery devel
opment plans along the Orissa coast.
FIGURE 1.7 The olive ridley, Lepidochelys olivacea.
There is some evidence that major “arribadas” are not permanent phenomena,
but rather they may wax and wane over cycles of at least several decades. Several
Mexican arribadas have disappeared in recent decades, and numbers are dropping
rapidly at the protected Nancite site in Costa Rica. Furthermore, the recent discovery
of a new site in Orissa, India, may correspond to the collapse of one of the two
previously documented arribadas along this coast. A former minor arribada in Suri
nam (the only one in the Atlantic) has now been decimated, presumably by a
combination of heavy egg collection for several decades and ongoing trawler mor
tality.
1.11.2,7 The Kemp's Ridley, Lepidochelys kempi
A 1947 film exists of an arribada of L. kempi estimated at 40,000 turtles at Rancho

Nuevo, Tamaulipas, Mexico. Apart from a few isolated cases, this was and is the
only known nesting site for this species. However, no subsequent data were available
from Rancho Nuevo until 1965, by which time the biggest arribada numbered less
than 5000 turtles, and despite beach protection this number continued to drop for
the next 20 years, by which time total nestings for the season numbered in the
hundreds only. Such nonresponse to protection is a classic indication of a population
for which recruitment has been nullified by near-total harvest of eggs, in this case
exacerbated by the intensity of shrimp trawling in the Gulf of Mexico (the primary
juvenile and subadult habitat and the only habitat of adult L. kempi) and, during at
least one episode in the late 1960s, beach slaughter of adults also (P. Burchfield,
personal communication). The nesting population reached a low point in the mid-
1980s, and has subsequently increased modestly, but steadily, possibly as a result
of the “head-starting” program initiated in 1978, about 10 years (thought to corre
spond to the approximate maturation time of the species) after the commencement
of the head-starting program. The species remains the rarest sea turtle in the world,
but the current apparent annual population increase may identify it more as “con
servation dependent” than actually “endangered.”
i |l ® i
FIGURE 1.8 The Kemp’s ridley, Lepidochelys kempi.
1.11.2.8 The Leatherback, Dermochelys conacea
None of the major nesting grounds of this species were discovered before the 1950s,
and many of them only in the 1960s or 1970s, and thus it is impossible to compare
contemporary population estimates with those of earlier in the century. Pritchard
(1982) noted recent discoveries of major nesting grounds in Mexico, and summarized
preceding discoveries that led to progressively increasing world population esti
mates.
The leatherback does not feature in international commerce, and its juvenile
stages (indeed, all stages between hatchling and adult) remain so cryptic that it is
FIGURE 1.9 The leatherback, Dermochelys coriacea.
unlikely that humans have any effect upon them. On the other hand, subsistence
take of eggs, and sometimes of nesting adults also, has been intense, especially in
the Eastern Pacific and Guyana, and while Asiatic nesting colonies (such as that in
Terengganu, Malaysia) are generally exploited for eggs rather than meat, this can
be equally devastating.
At present, the Atlantic colonies (especially in Trinidad, Suriname, and French
Guiana) appear to be reasonably secure and even increasing, as is the small nesting
colony in Natal, South Africa, and adjacent Mozambique. On the other hand, the
Terengganu colony has collapsed in recent years, and serious declines have been
documented in Pacific Mexico and Costa Rica, a result of the combination of beach
slaughter, egg collection, and serious incidental captures by fishing gear in the open
sea. Spotila et al. (in press) speculate that the species may be threatened with actual
extinction within the forseeable future, although Pritchard (in press) disagreed.
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The

CRC MARINE SCIENCE SERIES

Michael J. Kennish^ Ph.D.

The biology of sea turtles / edited by Peter L. Lutz and John A.

Visit the CRC Press Web site at www.crcpress.com

© 1997 CRC Press

Karen A. Bjorndal, Ph.D. Kenneth J. Lohmann, Ph.D.

Colin J. Limpus, Ph.D. David W. Owens, Ph.D.

Pamela T. Plotkin, Ph.D. James R. Spotila, Ph.D.

Blair E. Witherington, Ph.D.

Jeanette Wyneken, Ph.D.

1.2 MARINE TURTLES

1.3 HISTORICAL CLASSIFICATIONS OF MARINE

1.4 CLADISTIC PLACEMENT OF MARINE TURTLE

1.5 MARINE TURTLE FAMILIES

FIGURE 1.1 Cladogram “D”, Chelonidea.

1.6 FAMILIAL DEFINITIONS

A family of turtles characterized by an extensively roofed skull with well-developed

peripherals (often reduced to narrow bars in cheloniids). The plastron, primitively

1.7 DERMOCHELYID DIVERSITY

1.8 CHELONIID DIVERSITY

1.9 GENERIC DEFINITIONS OF LIVING CHELONIIDS

1.10 AFFINITIES OF RECENT CHELONIID SPECIES:

1. What subfamilies (or “Tribes”) should be recognized, and what is the

Deraniyagala (1939) recognized a superfamily, the Chelonioidae, comprising

Eretmochelys with Chelonia, and 15 justifying placement of Eretmochelys with

Systematists have been divided in recent years both in whether to recognize

1.11 SURVIVAL STATUS OF LIVING CHELONIIDS

1.11.2 B iologïcal S tatus

The actual, biological status of marine turtles is a topic of considerable complexity.

to a lower position once anthropogenic stresses (directed or incidental) become

1.11.2.1 The Green Turtle, Chelonia mydas

FIGURE 1.2 The green turtle, Chelonia mydas.

1.11.2.2 The Black Turtle, Chelonia agassizii

This species (possibly only a subspecies of C. mydas) is confined to the eastern

1.11.2.3 The Flatback, Natator depressus

FIGUKIC 1.3 "Hie black tuide, Chekmia agassizM.

FIGURE 1.4 The flatback, Natator depressus.

1.11.2.4 The Loggerhead, Caretta caretta

FIGURE 1.5 The loggerhead. Caretta caretta.

The small or medium-size populations in the Mediterranean, nesting primarily

1.11.2.5 The Hawksbill, Eretmochelys imbricata

The black turtle

FIGURE 1.6 The hawkshill, Eretmochelys imbricata.

1.11.2.6 The Olive Ridley^ Lepidochelys oUvacea

FIGURE 1.7 The olive ridley, Lepidochelys olivacea.

1.11.2,7 The Kemp's Ridley, Lepidochelys kempi

A 1947 film exists of an arribada of L. kempi estimated at 40,000 turtles at Rancho

FIGURE 1.8 The Kemp’s ridley, Lepidochelys kempi.

1.11.2.8 The Leatherback, Dermochelys conacea

FIGURE 1.9 The leatherback, Dermochelys coriacea.

Boulenger, G. A., Catalogue of the Chelonians, Rhynchocephalians, and Crocodiles in the

Mlynarski, M., Results of the Polish-Mongolian Palaeontological Expeditions — Part IV.

Population Genetics, Phylogeography, and Molecular Evolution

Reproduction In Sea Turtles

Orientation, Navigation, and Natal Beach Homing in Sea Turtles

Foraging Ecology and Nutrition of Sea Turtles

Age, Growth, and Population Dynamics

Hormones in the Life History of Sea Turtles

Salt, Water, and pH Balance in the Sea Turtle

Health Problems and Diseases of Sea Turtles

Human Impacts on Sea Turtle Survival

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