Richard Ellis Sea Dragons Predators of The Preh
Richard Ellis Sea Dragons Predators of The Preh
Richard Ellis Sea Dragons Predators of The Preh
RICHARD
ELLIS
2OOJ b y R i c h a r d L l l i s
All rights reserved
P u b l i s h e d b y the University Press o f K a n s a s
(Lawrence, Kansas 66049), which was organized
by the K a n s a s B o a r d of R e g e n t s and is o p e r a t e d
and funded by E m p o r i a State University,
Fort H a y s State University, Kansas State
University, Pittsburg State University,
the University of Kansas, and W i c h i t a
State University
Kllis, R i c h a r d .
S e a d r a g o n s : p r e d a t o r s o f t h e p r e h i s t o r i c o c e a n s / R i c h a r d 1:11
p. cm.
Includes bibliographical references and index.
ISBN 0 - 7 0 0 6 - 1 2 6 9 - 6 ( c l o t h : a l k . p a p e r )
1. M a r i n e reptiles, Fossil. 2. P a l e o n t o l o g y M e s o z o i c . I. T i t l e .
QE861.E45 2003
567-9' * 7 d c 2 i
2003006871
British L i b r a r y C a t a l o g u i n g - i n - P u b l i c a t i o n D a t a is available.
P r i n t e d i n the U n i t e d States o f A m e r i c a
1 0
Contents
Acknowledgments vii
Introduction: Isn't That the Loch Ness Monster? 1
T h e Marine Reptiles: A n Overview 1 7
T h e Ichthyosaurs 61
T h e Plesiosaurs 1 1 7
T h e Pliosaurs
T h e Mosasaurs
165
195
Acknowledgments
W h e n the world and I were somewhat younger, I started the research for a
hook that was going to he about the origin, evolution, and extinction of life in
the sea. I wanted to write about the formation of the earth, the earliest and
the later life-forms invertebrates to vertebrates, trilobites to a m m o n i t e s ,
lunghshes to whales. I was going to devote a certain amount of space to the
fate of those creatures that are no longer with us and those that still are, as
"living fossils." 1 thought I would also discuss the enigmatic disappearance of
the dinosaurs and the ancient marine reptiles. It soon became obvious that
those subjects were too many and too diverse to incorporate into a single
book, so in a fashion that has come to define my m o d u s operandi, I reduced
the scope of the project, kept some of the material I had written, and filed the
rest away for future use. In the original plan, one of the first subjects I tackled
was the marine reptiles, because they seemed so close in spirit and habitat to
other large vertebrates I had already dealt with sharks and whales and also
because, as far as I knew, there hadn't been a proper book written about these
neglected creatures since S a m u e l Williston's 1914 Water Reptiles Past and Present.
(In 1997, Academic Press published Ancient Marine Reptiles, but this was a
collection of disparate articles, and although they were all important, taken as
a whole, it was not the book I had in m i n d . )
N o t surprisingly, the material about ichthyosaurs, plesiosaurs, and m o s a saurs was so extensive that it soon became evident that these marvelous
creatures deserved more than a s u m m a r y chapter in a book about everything,
so they became the first casualties. ( A l s o put aside was the question of
whether there was water on M a r s or on Jupiter's moon Europa; a discussion
of the fascinating hut ridiculously wrongheaded idea that all the fossils ol
Archaeopteryx were forgeries; and the whole section on extinction past, present, and future which will become another b o o k . ) In 2001, Aauagenesis: The
Origin and Evolution of Life in the Sea, was published, and I still had a lot of neat
stuff left over. In a manner of speaking, this book was rescued from the scrap
heap.
thankful for c-mail, but they may not b e ) with exquisite patience were Dave
M a r t i l l of the University of Portsmouth (U.K.), Peter Doyle of the University of Greenwich (U.K.), Bill Sarjcant of the University of Saskatchewan,
Dale Russell of N o r t h Carolina State University, M i k e Caldwell of the
University of Alberta in Edmonton, Judy Massare of the State University of
N e w York at Brockport, L a r r y W i t m e r of the O h i o University College of
Osteopathic M e d i c i n e , R o b i n O'Keefe of the N e w York College of O s teopathic M e d i c i n e , M i c h a e l M a i s c h of Tubingen, N a t h a l i e Bardct of the
Natural H i s t o r y M u s e u m in Paris, Anne Schulp of the Natuurhistische
M u s e u m in Maastricht, Ben Kear of the South Australian M u s e u m in Adelaide, C a i t l i n Kicrnan, Kazuo Takahashi, Douglas Palmer, and Dan Varncr.
At a s y m p o s i u m in Lawrence, Kansas, on M o b y Dick in American art, I
met the editor of the accompanying book, Unpaitited to the Last. T h i s was M i k e
Briggs, editor-in-chief of the University Press of Kansas, and when he agreed
to publish this book, I dusted off the old hies about mosasaurs, ichthyosaurs,
and plcsiosaurs and began to do the drawings that I thought would help
reveal the long-hidden mysteries of the extinct marine reptiles. He edited this
book too, and our chance meeting over a discussion of the white whale has
produced a durable relationship of m u t u a l respect and admiration. As he has
done for the past 30 years, my agent Carl Brandt managed to place my
ungainly and overstuffed manuscript in the capable hands of someone who
could turn it into a proper book. Once again, Stephanie was along for the
ride, and as always, I am grateful for her support and loyalty. She keeps me
happy, and she also keeps me honest.
Introduction
Isn't That the Loch Ness Monster?
C o m e with me to the American M u s e u m of Natural H i s t o r y on N e w York
City's Central Park West. We walk up the broad stairs, d o m i n a t e d by the
equestrian
statue of T h e o d o r e
Roosevelt,
twenty-sixth
president of the
one. ( T h e " l i t t l e " Barosaurus is 32 feet long; the full-grown mother downstairs
measures 90 feet from nose to tail tip.) But keep going around the corner, and
here we are. L o o k up. T h e first thing you see hanging from the ceiling is
Dunkleosteus, a weird-looking, 20-foot-long armored fish with jaws that resemble overgrown staple removers. Wasn't there a gigantic shark jaw around here
somewhere? Oh yes, there it is, but when did it shrink? (It was rebuilt when
the curators realized that it had been m a d e half again as big as it actually was,
because the original fabricators made all the palm-sized teeth the same size
instead of m a k i n g the ones at the corners much smaller.) To the left is the
entrance to the library (probably the best natural history library in the
country, if not in the w o r l d ) , but at the moment, we're going to continue to
look up, as if bird-watching. A n d there, soaring high over the exhibit cases, is
probably the single most astonishing fossil in a m u s e u m filled with astonishing fossils. It has a long neck and a tiny head; a broad rib cage with a second
set of auxiliary ribs around the belly; a short tail ( m u c h shorter than its n e c k ) ;
and four broad fins, each with broadly flattened wrist bones and five long
"fingers." W h a t is the Loch Ness monster doing in the M u s e u m of Natural
History?
Better read the label to find out. It's k i n d of hard to find it, because the
skeleton is way up in the air, but here it is:
P L E S I O S A U R S
In this discussion and those that follow, it will become obvious that none of
these creatures has a c o m m o n name. We are used to referring to familiar
animals by their vernacular names, such as lion, tiger, fox, whale, h u m mingbird, jellyfish, and so on. T h e s e a n i m a l s also have scientific names, based
on a system developed by Carl von Linne ( C a r o l u s L i n n a e u s ) in the m i d eighteenth century. T h e lion is Panthera leo, the tiger is Pantbera tigris, the red fox
is Vulpes vulpes, the blue whale is Balaenoptera museulus, the ruby-throated hum-
hunting since 1933. Christian Spurling, WcthercH's stepson, revealed on his deathbed that lie
had participated in the manufacture and photographing of a foot-high model mounted on
a toy submarine, and they had coerced Dr. Wilson otherwise a man of impeccable
credentials to claim that he had taken the picture, when in fact it had been taken by
Wetherell (Langton 1 9 9 4 ) .
Skeletal
recon-
struction
of
of these names identifies the genus ( l i o n s and tigers are in the genus Panthera),
Plesiosaurus
dolichodeirus,
one ojthe
first
clasmosaurs
in
England.
jound
and the second identifies the species. In all cases, the genus (generic) name is
capitalized, and the species (specific) name is not. Lion, tiger, blue whale, and
so forth anY the names of these animals in English; in other languages, of
course, they are different. But whatever the language, the scientific name
remains the same. In a Polish, Chinese, Swedish, or Sanskrit discussion of the
lion, its scientific name still appears exactly as you see it here: Panthera leo.
T h e r e arc no equivalents of lions or tigers among the marine reptiles. Just
as with the terrestrial dinosaurs, the scientific name is the only one used.
Tyrannosaurus rex is sometimes shortened to T. rex, but it is still known only by
its scientific name. T h e same is true for Triceratops, Stegosaurus, Apatosaurus,
Veloeiraptor, and even Archaeopteryx. T h e names of the major groups of marine
reptiles can be rendered into English, as in ichthyosaurs, plcsiosaurs, and
mosasaurs, but each of these is also the generic name of a species within the
larger category, such as Ichthyosaurus communis, Plesiosaurus dolichodeirus, and Mosasaurus hoffmanni. S o m e of these names, like dolichodeirus, are more than a little
difficult to pronounce (it should be pronounced D O L - i k - o - D I E - r u s ) , but
because nobody ever refers to this species as "longncck" (the meaning of
dolichodeirus), whenever this species appears in print it is Plesiosaurus dolichodeirus.
In this introduction to the marine reptiles, you will encounter jawbreakers like
Ophthalmosaurus, Brachauchenius, and Pachycostasaurus, but most of the names are
somewhat easier and more comfortable on the tongue. T h e lack of common
names might even help in recognizing the most significant characteristic of all
the animals in this book: they are all extinct, and have been for m i l l i o n s of
years. We would like to become more familiar with them, but time and
nomenclature still remain formidable barriers.
As with many things palcontological, the evidence for the existence of longextinct creatures consists of bones. A n a t o m i s t s have given names to these
bones, and they are the same for living a n i m a l s as they are for extinct ones.
T h e bones in your arm below the elbow are the radius and ulna, and they are
the same although sometimes of greatly differing size and proportion for
whales, zebras, chipmunks, and dinosaurs. Because there are very few instances in which anything but bones is preserved and in those cases, rarely
completely those who describe extinct a n i m a l s often limit themselves to
detailed descriptions of the bones. If, for example, one finds a fossilized
ichthyosaur with an upper jaw longer than that of another known fossil, the
long-jawed ichthyosaur might be described as a new species. It m i g h t also be a
juvenile as opposed to an adult, but other measurements can confirm its
similarity to or difference from other known specimens.
M a n y of the descriptions of the creatures in this book all of which are
extinct, and all of which arc known only from fossils consist primarily of
osteological terminology. ( O s t e o l o g y is the study of bones.) Even the size of
the eye, so critical to the differentiation of various ichthyosaur genera, relies
largely on the circle of bony plates in the eye socket known as the sclerotic
ring. At least part of the behavior of a large-eyed animal can be postulated
from the size of the sclerotic ring, and although we might suggest that such a
creature hunted at night or in reduced light circumstances, we can only guess
as to what it hunted. ( S o m e t i m e s , remnants of its last meal are fossilized too;
in some cases, squid beaks or sucker hooks have been found in the fossilized
predator's stomach.) T h e ability to separate one species from another d e pends on these detailed descriptions, and when one fossil is compared with
another and found to be different in its particulars, the result m i g h t be a new
species. Comparative anatomy therefore is one of the cornerstones of paleontology, but it often results in complex technical descriptions not easily understood by the nonspecialist. Here, for example, is Edward Drinker Cope's
(1868a) description of a plcsiosaur fossil that had been shipped to him from
Kansas:
T h e species represented a genus differing in important features from
Plesiosaurus and its near allies. T h e s e were the absence of diapophyses on
the caudal vertebrae, and the presence of inferiorly directed plate-like
parapophyscs which took the place of the usual chevron bones, in the same
position; also in the presence of chevron-like bones on the inferior surfaces
of the cervical vertebrae; further in some details of the scapular and pelvic
arches. T h e diapophyses of the dorsal vertebrae originated from the centrum, and not from the neural arch.
C o p e was reading the vertebrae backward, which resulted in his reconstructing the skeleton with the skull at the wrong end, but the point is the same:
"the presence of inferiorly directed plate-like parapophyscs" docs not help the
layperson understand what Elasuwsaurus platyurus looked like. Most descriptions of fossils are like this, and only the most creative of paleontologists, or
those attempting to make particular points about l i m b structure and movement or tooth structure and prey items, will extrapolate from the bones to the
lifestyle. Because we are not all trained paleontologists, we would like to
learn more about a given species than we can discover from the absence of
diapophyses.
Although there are no " r u l e s " governing the form of a scientific paper,
most of them follow a recognized pattern, generally consisting of abstract,
introduction, discussion, conclusion, and references. In a paleontological
discussion, there is usually a section describing the location of the fossil find,
its condition, and its eventual disposition; if possible, an attempt is made to
place it in a recognizable phylogenetic category, such as ichthyosauridac or
m o s a s a u n d a e , so the reader will know the general nature of the fossil. (It is
not always easy to place, say, a single tooth in a known category, and there
have been instances when the description of the tooth was accurate but its
designation was not.) Such organization is possible for scientific papers, but it
is considerably more difficult with books. T h i s book has certainly been
organized into broad categories (ichthyosaurs, plesiosaurs, pliosaurs, mosasaurs), but the very nature of paleontology, with new specimens being un-
earthed or descriptions of old ones being revised in the literature, makes for
an uneasy chronology. Does the writer talk about the sequence of discovery or
the geological sequence of the specimens themselves? Is the "earliest" ichthyosaur the oldest, or the first one found by fossil hunters? T h e first discovery of a fossil mosasaurin fact, the first discovery of a fossil marine reptile
of any k i n d occurred in a Belgian limestone mine in 1780. Later christened
Mosasaurus hoffmanni, it was found to be one of the last of the mosasaurs;
therefore, depending on who's doing the structuring, the story of the mosasaurs can cither begin at the end or end at the beginning.
M a n y early paleontologists a t t e m p t e d to reconstruct the lifestyle of the
marine reptiles or the dinosaurs, but because this exercise was speculative,
often the safest thing to do was to describe the bones and leave the lifestyle to
someone else. It is true that much can be learned from the size and shape of
the bones and the muscle attachments, and from this evidence it can be
ascertained how the animal might have moved, but what it did when it got
there is often an enigma. M u c h can also be deduced from teeth. Just like
today's animals, extinct large animals with big, sharp teeth were probably
aggressive killers, and it is not difficult to imagine that a big marine lizard
with teeth like a crocodile or a killer whale might have behaved in a similar
fashion to these powerful predators. A n i m a l s with flattened teeth that look
suitable for grinding were probably plant or shellfish eaters. Osteological
descriptions are critical to understanding the relationships of extinct animals,
but because we cannot observe them in action, we can only guess as to how
the animals swam or hunted or gave birth. W h e n e v e r possible, therefore, I
describe the animal's size, teeth, flippers, and tail ( o r where the fossil was
found and by w h o m ) and refrain from differentiating various species by the
relative size and shape of the shoulder blade or pelvic girdle.
In many cases, there is enough fossil evidence to allow a fairly accurate
reconstruction of the animal's size and shape, but except for the obvious
eves are for seeing, teeth are for biting or tearing, backbones arc for support
conclusions can rarely be drawn about how the animal actually used this
equipment. As will be seen, the existence of four flippers in plesiosaurs has
presented a virtually unsolvable question of how the flippers were used to
propel the animals through the water, but we have a pretty good idea of the
prey subdued by the e n o r m o u s teeth of the giant pliosaurs. M o s t ichthyosaurs had a downward tailbcnd at the end of the vertebral column, which
suggests that their s w i m m i n g was powered by flexions of the lower lobe, so
the comparison to the tail of a shark where the vertebral column extends
into the upper lobe is obvious. We know what the tails of living sharks look
like, but in only a few cases in which the outline of the entire animal was
preserved do we know what the upper lobe of an ichthyosaur's tail looked like.
T h e mosasaurs had scales, not unlike those of a snake, and at least some
ichthyosaurs had smooth skin like that of whales and dolphins ( L i n g h a m S o l i a r 2001); otherwise, with so few preserved fossilized impressions, we
do not know whether the rest of them particularly the plesiosaurs were
smooth-skinned or had scales, ridges, l u m p s , or bumps. And there is one
thing that we know nothing about: we have no idea what color the marine
reptiles were.
In order to re-create them, I could have made them monochromatic (I was,
after all, working in pen and i n k ) , but this seemed to detract from their
vitality. We know that many living reptiles are brightly colored think of
snakes and lizards so there is no reason to assume that the reptiles that lived
100 m i l l i o n s years ago were drab and colorless. Because these reptiles lived in
the water, does that mean that they would be only countershaded, like many
fishes dark above and light below? Of course not. Fishes come in all the
colors of the rainbow, and for g o o d measure, some of them even light up.
Well, then, aren't whales dull and monochromatic? N o t on your tintype.
Killer whales are spectacularly patterned in black and white; various dolphins
sport elaborate haberdashery; and the fin whale, with its complicated a s y m metrical pattern of swoops and swirls, may be the most intricately colored
animal on Earth. In my drawings, I made some of the reptiles plain, and I
patterned some of them with stripes, spots, and countershadings. All these
color schemes are imaginary, designed (I h o p e ) to breathe life into long-dead
ichthyosaurs, plesiosaurs, and mosasaurs.
T h e fossil record is tantalizingly incomplete. T h e r e are any number of
creatures that lived on Earth for which we have found no evidence whatsoever,
so paleontologists have had to make do with the comparatively small number
of fossils found and extrapolate from there. In Atlas of the Prehistoric World,
Douglas Palmer wrote, "there should be in the order of 500 million fossil
species buried in the stratigraphic record. So far, paleontologists have described only a few hundred thousand fossil species. At less than 0.01 percent,
this represents a very small sample of the estimated total." (Palmer says that
real numbers are not available and his figure is a "guesstimate," but it is
probably in the right range.) In 1994, David R a u p summarized the fossil
record for dinosaurs as follows:
T h e dinosaur fossil record illustrates some of the more severe sampling
problems. According to a review by Dodson, 336 of the named species of
dinosaurs are taxonomically valid. Of these, 50% are known only from a
single specimen, and about 80% are based on incomplete skeletons. T h e
336 species are grouped into 285 genera, and of these, 7 2 % have been found
in the rock formations where they were first discovered, and 7 8 % have been
found in only one country. T h e s e numbers are astonishing if viewed as if
the data were complete.
In many instances, a single tooth, bone, or bone fragment has been found,
and because there is no other possible explanation, the paleontologist identifies the animal that originally owned these bones and declares that it once
lived ( o r d i e d ) here. H o w do they know? C o m p a r i s o n with specimens d e scribed in books and journals and those seen in museum collections, experience, and, of course, location. A great many species have been described from
limited fragments of evidence, and the discovery of a complete or even
partially complete specimen is a rare occurrence in paleontology. T h e r e are
some notable exceptions where fossils are particularly plentiful, and they are
occasionally even fairly complete. T h e s e special sites include the Burgess
Shale in British C o l u m b i a , H o l z m a d e n and Solnhofen in Germany, the seaside cliffs of Dorset in England, certain areas of the Gobi Desert, the H e l l
Creek Formation of M o n t a n a and the Dakotas, the Bear Gulch Formation in
M o n t a n a , the Niobrara C h a l k F o r m a t i o n s of Kansas, and the Yixian Formation of the Liaoning province of C h i n a . Even when the fossils are relatively
complete, an e n o r m o u s amount of work is required to extract them and
prepare them for study or exhibition.
Consider the fossils unearthed by N e w Zealand's "Dragon Lady," Joan
missing lineages, or even missing t a x a ) , there are still many ways that good,
testable hypotheses can be developed in studying the fossil record. Of necessity, the evidence collected is usually more indirect and circumstantial, but
that does not make it less worthy than direct evidence, if used properly.
Because paleontologists do not have the o p p o r t u n i t y to observe the living
subjects of their studies, their hypotheses must be tested by techniques of
comparative biology, based on a thorough, detailed, and broad knowledge of
living animals. T h e r e are no living ichthyosaurs, but they shared certain
characteristics with sharks and dolphins, and comparisons with the living
animals have given us great insights into the m o d u s V i v e n d i o f the fish lizards.
Mosasaurs resemble varanid lizards in some respects and crocodilians in
others, and comparative studies have enabled paleontologists to make numerous assumptions about the lives and phvlogenv of the great seagoing
lizards. Unfortunately, there arc few living creatures that resemble plesiosaurs
(only today's sea turtles propel themselves with four flippers), and so much
about their lives is still a mystery. T h e absence of living models, however, does
not preclude creative analysis of the fossils. T h e r e is a substantial body of
literature devoted to the locomotion of plesiosaurs, all based on the shape,
structure, and relationship of the bony elements. Indeed, one does not have to
compare fossil structures to analogous structures in living animals; the hydrodynamic capabilities of plcsiosaur flippers have been compared to the wings
of birds, bats, and even airplanes (O'Keefe 1 0 0 1 c ) .
T h r o u g h o u t this discussion of the marine reptiles, I cite the various chronological periods, which have been named by geologists and paleontologists so
that they would have a consistent timetable with which to associate particular
fossil faunas. T h e span under discussion here is generally known as the
M e s o z o i c era, which lasted from 248 million to 6$ million years ago. T h e
M c s o z o i c is further broken down into three large periods, the Triassic (248 to
209 million years a g o ) , the Jurassic (208 to 144 million years a g o ) , and the
Cretaceous (144 to 65 million years a g o ) . T h e s e have been further compartmentalized into smaller, tighter groups; the late Cretaceous, for example, is
subdivided into the Coniacian age (89.9 million years a g o ) , the Santonian age
(85.8 million years a g o ) , the C a m p a n i a n age (83.5 m i l l i o n years a g o ) , and the
familiar with the comparison of the history of life on Earth to a 24-hour day,
in which humans have been around for only the last few seconds, but such a
construct diminishes the actual passage of time, probably in the ever-present
interest of anthropocentrism the belief that the world revolves around us
and that we can understand things only in human terms. But human beings, as
we know us, have been around for 100,000 years, a fleeting one-tenth of a
million. T h e time of the mosasaurs, often described as "only" 25 million
years, is 250 times greater than the total experience of Homo sapiens, from the
moment he (or she) picked up the first rock and shaped it into an ax head to
the moment you are reading these words. T h e ichthyosaurs lasted four times
longer than the mosasaurs, so their time on earth was a thousand times longer
than ours. To grasp the pace of evolution, we don't need to speed up the film,
we need to slow it down. We must not be misled by the idea that a million
years is a mere blink of the eye. T h a t is the case only in geological terms; the
planet is believed to be 4.5 billion years old, but a million years is a very long
time indeed. If a human generation is 20 years, then 5,000 generations have
passed in the entire history of H. sapiens, and if we were to last a million years
a highly unlikely scenario 50,000 generations would pass. If a generation of
ichthyosaurs was also 20 years, then there were 7.5 million generations of fish
lizards in their 150-milIion-year history. Evolution is a slow process, but
during their "reign," the dinosaurs diversified into hundreds of different
species, grew to enormous sizes, and even sprouted wings and feathers. Various ancestral reptiles took to the water, and through an inexorably slow and
gradual process comparatively speaking, glaciers move at the speed of
bullets became the aquatic reptiles that will be visited here.
T h e aquatic reptiles are all believed to have descended from terrestrial
forebears, but those forebears were descended from animals that lived in the
water. T h e first terrestrial tetrapods of the late Devonian period (circa 354
million years ago), such as Acanthostega and lehthyostega, emerged from the water
with their limbs modified to walk on land. Recent discoveries ( C o a t e s and
Clack 1990) indicate that these early tetrapods had eight digits on the forelimbs and seven on the hind. T h i s plan d i d not perdure, and the five-finger
arrangement d o m i n a t e d the future of reptiles and m a m m a l s . (Stephen J.
Gould wrote an essay on seven- and eight-fingered tetrapods, suggesting that
the
ichthyosaurs) arc known unequivocally to have given birth to live young in the
water, but the evidence is less convincing for the plesiosaurs and mosasaurs;
and they were all descended from terrestrial reptilian ancestors. Some of them
were contemporaries in time and place, but the ichthyosaurs finally went
extinct at the Cenomanian-Turonian boundary, which was 93.5 million years
ago, or 25 m i l l i o n years before the demise of the last of the plesiosaurs and
mosasaurs. T h e final extinction of the plesiosaurs and mosasaurs is thought
to be somehow connected to the event that took out the nonavian dinosaurs
65 m i l l i o n years ago, but how an asteroid impact and its consequences eliminated some of the seagoing reptiles while sparing the turtles and crocodiles is
not clear.
Despite the apparent similarities in habitat and lifestyle, however, the three
major groups of marine reptiles were quite different and were not closely
related. T h e ichthyosaurs were more or less dolphin-shaped, but they had
four flippers to the d o l p h i n s ' two and a vertical tail fin where that of the
* In
question. I low can .1 bird be a reptile? "( Heady we have a decidedly different Reptilia twin
the traditional motley crew of crawling, scaly, nonmammal, nonbird, nonamphibian creatures that most ol us think of when we think of reptiles. If it is true that crocodiles and
birds are more closely related to each other than cither is to snakes and lizards, then a
monophyletic group that includes snakes, lizards, and crocodiles must also include birds.
I he implication of calling a bird a reptile is that birds share the derived characters of
Reptilia, as well as having unique characters of their own."
none were mentioned by Linnaeus, because none of them had been recognized before he died in 1778.
Because Linnaeus was working in the m i d d l e of the eighteenth century, he
was able to include some newly discovered beasts in his classification, but he
firmly believed that all known species were unchanging creations of God, who
had chosen to arrange things so that man resided at the top of the hierarchy.
W h a l e s and dolphins had been known since they demonstrated the unfortunate inclination to beach themselves. Two thousand years ago, Aristotle
wrote, "It is not known for what reason they run themselves aground on dry
land; at all events, it is said that they do so at times, and for no obvious
reason." U n t i l whalers and other seafarers took to the sea, the only cetaceans
that could be known were those that washed ashore. T h e discovery of fossils,
known since ancient Greece, suggested that there were some life-forms that
were no longer with us, but for European and American minds, this meant
only that the Great Flood, identified in the Bible, had drowned those that
N o a h hadn't loaded onto the ark. ( H o w the fishes, whales, and dolphins
managed to board the ark was never explained; maybe they swam along in its
wake.) As humankind's horizons widened, animals that Linnaeus never knew
of began to appear. In N o r t h America, there were raccoons, pronghorns, and
m o u n t a i n lions; South America had weird and wonderful monkeys with
prehensile tails, sloths, anteaters, llamas, and a r m a d i l l o s ; and there was an
entire continent in the southern sea that was populated by the strangest fauna
of all: kangaroos, wallabies, koalas, and w o m b a t s mammals that raised their
young in a pouch. T h e "dark continent" of Africa had giraffes, water buffaloes, zebras, baboons, chimpanzees, and gorillas, not one of which was
mentioned in the story of the biblical flood.
N o n e of these animals, no matter how bizarre, would astonish the scientist
and the layperson the way the great fossil reptiles did. After all, deer and
antelopes are relatively familiar; llamas sort of look like camels, and even the
fantastic fauna of Australia fit albeit roughly into a range of sizes and
shapes that would hardly cause people to question their fundamental beliefs
about life on Earth. But here we had evidence of 40-foot-Iong lizards with
huge teeth; dolphin-like creatures with long snouts and four legs; and, most
wondrous of all, a long-necked something with a tiny head and what looked
like flippers! Everything people thought they knew about the living world was
being stood on its head. T h e r e was nothing remotely resembling a comprehensible framework that could enclose these fabulous creatures. A whole new
wav ol looking at life would have to be developed and quickly. Fortunately,
there were men like Baron Cuvier, Geoffroy S a i n t - H i l a i r e , and the Chevalier
de Lamarck; Dean W i l l i a m Daniel Conybeare and Professor W i l l i a m Buckland; Richard Owen and H e n r y De la Beche; Carolus Linnaeus and Charles
Lyell; and, ultimately, the men who would furnish the system that could
incorporate the lizards, the llamas, the lions, and the kangaroos Charles
Darwin and Alfred Russcl Wallace.
Like the first of the dinosaurs, the first of the great sea reptiles was found in
England. Although the British were the first to find and publish descriptions
of the extinct marine and terrestrial reptiles, fossils would soon be appearing
in other European countries, in N o r t h America, and eventually on every
continent. In 1719, Dr. W i l l i a m S t u k e l y was apprised by R o b e r t Darwin
(Charles's grandfather) of a "human Sceleton (as it was then t h o u g h t ) " that
had been found in the bluestone quarries of N o t t i n g h a m s h i r e . Stukely examined the specimen and realized that "it cannot be reckoned H u m a n , but seems
to be a Crocodile or Porpoise." ( T h a t there were "no less than Eleven Joints of
the Tail" probably helped in his d i a g n o s i s . ) In his discussion of the "sceleton,"
he wrote: " W h a t Creature this has been, for want of a N a t u r a l H i s t o r y of
Scclctons, well worthy of the Endeavors of this society, we cannot possible
determine; but generally find the like to be a m p h i b i o u s or marine Animals."
Stukely s was the earliest authenticated reference to a plcsiosaur, but the name
hadn't been invented yet. M o r e than a century would pass before Conybeare
and De la Beche would coin the term in their 1824 discussion of Plesiosaurus
dolichodeirus, the long-necked near-lizard. (Stukely's "sceleton," now properly
identified as a plcsiosaur, is on exhibit in the Natural H i s t o r y M u s e u m ,
I .ondon.
In 1818, W i l l i a m Buckland, a professor of geology at Oxford, was shown
the bones of a huge carnivorous beast, and when he couldn't identify the
original owner, he sent them to Baron Cuvier, the w o r l d s foremost authority
on fossils and anatomy, who helped Buckland see that they had indeed come
from a giant reptile. Six years later, Buckland published "Notice on the
Megalosaurus, or the great fossil lizard of Stoncsficld." He did not call it a
dinosaur, because the term would not appear until Richard Owen introduced
it in 1842. He had the fossil lower jaw ( w i t h teeth in place), several vertebrae,
some fragments of the pelvis, and the shoulder bones of large, unknown
a n i m a l s that he had been collecting for about a decade from the Stoncsficld
quarries near Oxford. In his 1824 paper, Buckland identified it as a species of
giant extinct lizard (Megalosaurus means "great l i z a r d " ) that exceeded 40 feet in
length and had a bulk equal to that of a large elephant.
In 1821, while walking through T i l g a t e Forest in Sussex, M a r y Ann M a n tell found a toothlikc fossil that she gave to her husband, Dr. Gideon M a n tell, who went on to become a compulsive collector of fossils ( M a r y Ann left
him because there was no more room in their house, among other reasons).
M a n t c l l felt that the tooth did not belong to a crocodile or an ichthyosaur,*
so he sent it to Cuvier, who identified it as belonging to an extinct form of
rhinoceros. But when M a n t e l l compared the tooth with that of an iguana, he
realized that the tooth his wife had found was almost a replica of the tooth
of the living lizard, but 20 times larger. He named it Iguanodon ("iguanat o o t h " ) , and although it was in fact the first dinosaur ever described, it
entered the literature only as the " N e w l y discovered Fossil Reptile from the
sandstone of the T i l g a t e Forest in Sussex." Cuvier rescinded his misidentification in a letter to M a n t c l l in in 1824. Buckland and M a n t e l l both believed
that their creatures were ( o r had been) giant lizards, and the parade of the
dinosaurs had begun.
T h e seagoing reptiles were actually discovered a couple of years before the
land-dwelling dinosaurs and were therefore the first prehistoric animals to
* According to Chris McGowan ( 2 0 0 1 ) , "This is a wonderful story of the discovery of one
of the most important fossils of the time. Unfortunately, it is without foundation, and
Mantell himself was largely to blame for this. Not only did he fail to record at the time how
the unique tooth was discovered, but he also gave somewhat diflerent accounts of the find
after the fact. . . . Dennis Dean, an authority on Mantell, is of the opinion that the first
tooth was probably supplied to Mantcll by Mr. Lcncy, the quarryman at Cuckfield. . . .
Regardless of the provenance of the first tooth, it must have astonished and baffled Mantell
because it was so entirely different from anything he had ever seen before."
T h e s e animals attacked one another with inconceivable fury. Such a combat was never seen before by m o r t a l eyes, and to us who did see it, it appeared
more like the phantasmagoric creation of a dream than anything else. T h e y
raised m o u n t a i n s of water, which dashed in spray over the raft, already
tossed to and fro by the waves. Twenty times we seemed on the point of
being upset and hurled headlong into the waves. H i d e o u s hisses appeared
to shake the g l o o m y granite roof of that m i g h t y cavern hisses which
carried terror to our hearts. T h e awful combatants held each other in a
tight embrace. I could not m a k e out one from the other. Still the combat
could not last forever; and woe unto us, whichsoever became the victor.
O n e hour, two hours, three hours passed away, without any decisive result.
In the end, the Ichthyosaurus triumphs and "returns to his m i g h t y cavern
under the sea to rest." It is obvious that Verne had read some of the contemporaneous descriptions of plesiosaurs and ichthyosaurs and had seen some of
the more spectacular illustrations, perhaps those in Louis Figuier's 1863 La
Terre avant le Deluge (Earth before the Deluge).* Figuier's book includes illustrations by Edouard R i o u , one of which depicts an ichthyosaur confronting a
plesiosaur, which M a r t i n R u d w i c k calls "a visual cliche." T h e discovery of
fossils of gigantic marine reptiles clearly captured the imagination of the
public. T h e ichthyosaurs and plesiosaurs were bigger, fiercer, and more terrifying than any puny crocodiles, and now it was up to the scientists to
explain where they had come from and, Jules Verne notwithstanding, what
had happened to them.
* As a novelist, it was Verne's prerogative to exaggerate the size and ferocity of the saurians,
but what Figuier actually wrote ( i n Rudwick's 1 9 9 2 translation) was: "we bring together
these two great marine reptiles of the Lias, the Ichthyosaur and the Plesiosaur. Cuvier says
of the Plesiosaurus 'that it presents the most monstrous assemblage of characteristics that
has been met with among the races of the ancient world.' It is not necessary to take this
expression literally; there are no monsters in nature; the laws of organization are never
positively infringed; and it is more accordant with the general perfection of creation to see
in an organization so special, m a structure which differs so notably from that of animals of
our days, the simple augmentation of a type, and sometimes also the beginning and
successive perfecting of these beings."
was m a k i n g it more and more unlikely.* After comparing the skulls of the
Indian and African elephants, Cuvier announced ("with remarkable selfassurance some m i g h t term it arrogance," wrote R u d w i c k ) that they were
different species and, furthermore, that the mastodon, recently unearthed in
Ohio, was distinct from either of the two living elephants. He believed that
the m a s t o d o n was extinct, as were the previous owners of many fossil animal
parts, but he was unable to determine why so m a n y living species had survived
the catastrophes while the extinct species (especes perdus) had not. Cuvier wrote
(translated in R u d w i c k 1972), "the most important question being to discover
if the species that then existed have been entirely destroyed, or if they have
merely been modified in their form, or s i m p l y transported from one climate
into another."
In 1808, Cuvier was shown the fossilized jaws of an e n o r m o u s reptile that
had been found in a limestone m i n e in M a a s t r i c h t in what is now the
Netherlands. T h e jaws looked something like those of a modern monitor
lizard, but they were more than 3 feet long. T h e jaws would later be described
by Conybeare as belonging to the first mosasaur ( n a m e d for the M e u s c River
near which they were found), a gigantic marine lizard that was also, mercifully,
extinct. Cuvier's ability to reconstruct entire animals from a single bone
meant that any unexplainable fossils would be brought to him for identification. From a small scrap, he was able not only to identify the animal it came
from but also to postulate its way of life. It was not long before he was shown
the bones of a reptile that appeared to have wings, which he named Pterodactyle, m e a n i n g "winged fingers." Cuvier realized that some animal species had
* But not impossible. During the twentieth century, when most people would assume that
all the large animals had already been discovered, the okapi was found in central Africa in
1 9 0 1 , the coelacanth off South Africa in 1 9 3 8 , and the mcgamouth shark in Hawaii in 1 9 7 5 . In
1 9 9 3 , the saola or Vu Quang ox (Pseudoryx nghetinensis), a previously unknown deer-sized
animal with horns like an antelope, was discovered in a mostly unexplored ram forest on the
Vietnam-Laos border. In 1 9 9 7 , scientists in Myanmar (formerly Burma) found the world's
smallest deer, Muntiacus putaoensis, about the size of a large beagle and half the size of the
smallest deer previously known. And in 1 9 9 9 , Pitman et al. identified Iniopacetus pacifitus, a
new species of bottlcnose whale from the tropical Indopacific a 25-foot-Iong cetacean
whose existence was suspected but unconfirmed.
disappeared at least from some parts of the world which meant that they
had been destroyed, had somehow t u r n e d into something else, or had wandered far from the spot where the fossils had been found and were still alive
somewhere. It would be another 40 years before Charles D a r w i n would
suggest that extinction was somehow related to evolution, but Cuvier proposed periodic mass extinctions caused by catastrophes like the biblical deluge, i le believed thai the unrecognizable forms appeared after each Hood and
that fossils were remnants of the most recent previous creation. ( H e would go
to his death, however, proclaiming that there would never be a h u m a n fossil;
he declaimed emphatically, I.'hommefossile n'existe pas!) For hundreds of years,
educated humankind had believed in the "Great C h a i n of Being" (Scala Naturae), a hierarchical arrangement of every living thing in the universe. T h e r e
were "base" metals like lead, and "noble" ones like gold and silver. Given the
authors of the list, it is not surprising to find man at the top, just below God
and the angels. T h e chain progressed incluctably upward, from the "lower"
animals such as insects and w o r m s to the "higher" animals, the birds and
mammals.
behind the orbit have been further separated into two principal groups
the lepidosaurs ("scaly reptiles"), which includes the living snakes and the
lizards (including the extinct mosasaurs), and the archosaurs ("ruling rept i l e s " ) , which includes crocodiles, dinosaurs, and flying reptiles. Although the
nothosaurs, placodonts, plesiosaurs, and ichthyosaurs had only a single skull
opening, they arc classified as diapsids, but of unknown relationships.
T h e first reptiles lived on land, as d i d the ancestors of the living crocodiles,
lizards, snakes, and tuataras. T h e s e proto-reptilcs were small and light-boned,
not unlike most lizards today, probably because smaller bodies were easier to
maintain on the small grubs, insects, and w o r m s that could be found on the
floor of the Carboniferous forests, and also because small bodies are easier to
heat up in whatever sunlight managed to break through the canopy. Richard
Cowen (2000) believes that the first reptiles evolved "cither on the rivcrbanks
or in the canopy ecosystem of the Early Carboniferous, not on the forest
floor," and may have even lived in the hollowed-out stumps of tree ferns.
However they accomplished it, Carboniferous reptilomorphs like Westlothiana
or Hylonomus were among the first animals on Earth to lay amniotic eggs, in
which the embryo develops in a leathery shell, instead of the jelly-covered
eggs of amphibians and fishes. U n l i k e typical amphibian and fish eggs, which
have to be laid in water, reptile eggs must be fertilized before they arc laid, and
they must be laid on land. T h e shell prevents the embryo from d r y i n g out,
and the developing reptile emerges as a miniature adult only when it is
competent to survive on its own. ( M o s t amphibians pass through an aquatic
larval phase, like the tadpole stage in frogs, before they come onto l a n d . ) All
the future amniotes (later reptiles, birds, and m a m m a l s ) developed because of
this innovation. But the reptiles, which are characterized by a scaly integument, never developed the ability to heat themselves mctabolically and were,
for the most part, dependent on ambient temperatures. ( W e don't really know
about the terrestrial dinosaurs, which were reptiles. But modern birds, which
are their descendants, are certainly w a r m - b l o o d e d . ) It is likely, however,
that some of the marine reptiles, particularly the ichthyosaurs, were w a r m blooded, as were some of the dinosaurs. S o m e of the marine reptiles even
"advanced" beyond the limitations of the amniotic egg, to the point where
they became viviparous, giving birth to live young underwater, the way whales
and d o l p h i n s do today.*
Reptiles evolved on land, but after their dispersal to various terrestrial
habitats, some returned to the sea. In geological time, this turnaround occurred soon after the conquest of the land, because there are records of
anapsids known as mcsosaurs ( " m i d d l e l i z a r d " ) that evidently achieved an
aquatic existence as soon as the early Permian, almost 300 million years ago.
Mesosaurus, the first known marine reptile, was a 3-foot-long, fully aquatic
lizard-like animal that propelled itself underwater with its elongated tail and
webbed hind feet. S o m e thought that its elongated jaws, equipped with fine,
sharp teeth that were too delicate for snagging fish, might have formed a sieve
to strain plankton from the water, but m o d e r n analyses render this an unlikely
explanation. T h e mcsosaurs died out about 250 million years ago and evidently left no direct descendants. Because mesosaur fossils have been found in
southern Africa and eastern S o u t h America, a good case can be made for
continental drift. Since Mesosaurus could not have s w u m across the ocean from
one continent to the other, its presence on both sides of what is now the
South Atlantic supports the notion that the two land masses were once
joined.
As S a m u e l W i l l i s t o n wrote in 1914, "Were there no turtles living we should
look upon the fossil forms as the strangest of all vertebrate animals animals
which had developed the strange habit of concealing themselves inside their
rilis. lor that is literally what the turtles do." If it wasn't for the 250 species of
turtles living today, trundling around on land and s w i m m i n g in the sea or in
lakes and ponds, these animals encased in mobile homes could easilv be
* Giving birth to live young (viviparity) may not be the "advance" that us live-bearers think
it is. In a letter to me, Mike Caldwell wrote, "Many groups of living squamates are
viviparous/ovoviviparous while closely related forms (sister taxa) are egg-layers (e.g.,
pythons lay eggs, boas give live birth). Which is more advanced? I don't know and couldn't
begin to guess. The problem intensifies as you go up the tree many vipers lay eggs while
crotalids (rattlesnakes) are live bearers. Between boas and rattlesnakes there are thousands of
species and millions of years. W h i c h is the advanced condition? Again I don't know. I do
not think the amniotic egg and shell is a limitation. Ichthyosaurs and mosasauroids simply
bypass the egg shell and retain the embryo in the body as do boas and rattlesnakes."
Only 3 feet
long,
Mesosaurus is
thought to he one of
the first reptiles to
return to the water.
viewed as bizarre evolutionary experiments that were ordained for failure. Sea
turtles were much more varied and diverse than they are today, but some of
would have
the earliest turtles, known from the late Triassic (200 million years a g o ) , had a
shell and an anapsid skull a solid block of bone, with no openings for jaw
water,
muscles. T h e first known turtle was Proganochelys quenstedi, which had a fully
developed shell and a turtle-like skull and beak, but it also had several
primitive features not found in turtles today, including tiny teeth on its palate,
a clavicle, and a simple car. T h e early turtles were unable to w i t h d r a w their
heads or legs into their shells, but by the m i d d l e of the Jurassic, turtles had
split into the two main groups of turtles found today, the side-nccked turtles
(plcurodires) and the arch-necked turtles ( c r y p t o d i r e s ) . T h e y probably had
terrestrial ancestors, but the sequence of descent is not evident.*
' Although this i s the traditional v i e w , Ricppcl and Reisz presented a paper in 1 9 9 9 in which
they argued that the opposite conclusion was possible: "Within Amniota, the turtle bodyplan is highly derived (autapomorphic) which results in functional constraints that are
indicative of an aquatic origin for turtles. The most important of these traits is the
helped
hut exactly
It lived
million
development of a carapax and plastron, which results in the reorganization of the paraxial
mesoderm in the trunk region. The consequences of this ontogenetic repatterning affects
locomotion and respiration in profound ways, necessitating structural and functional
changes that are more easily achieved in an aquatic rather than a terrestrial environment. An
aquatic origin of turtles is also suggested by the earliest appearance of the cladc in the
Middle Triassic Muschelkalk of Germany, an appearance that matches quite closely the
time of the origin of the turtle clade suggested by the molecular data."
Although
"tortoise-
30-inch-long
hawksbill
turtle
(Eretmochelys
imbricata) is still
hunted
its
throughout
worldwide
tropical habitat
its
beautifully
mottled
shell.
for
loggerhead is a
The
turtle
C a i v u . i caret la
has
a proportionally
between
400
zoo
pounds.
T h e smallest of the sea turtles are the ridlcys; the 2-foot-Iong Kemp's ridley
(I.epidochelys kempiif breeds only in the Gulf of M e x i c o , and the slightly larger
Pacific ridley (L. olivacea) is found in the Indian and Pacific Oceans. Kemp's
ridley is seriously endangered, and there m a y be no more than 1,500 left in the
world. T h e n o n m i g r a t o r y
flatback
northern Australia, is the only species of sea turtle that is not considered
threatened or endangered.
After the mesosaurs, the earliest known marine reptiles arc the nothosaurs,
which had slender bodies, long necks and tails, and webbed feet. T h e nominal
genus (Nothosaurus) had a long, narrow skull with splayed teeth that intcrmeshed when the mouth was shut, or, as R i e p p e l (2001b) put it, "As in all
species of Nothosaurus, the anteriormost two fangs erupt from their respective premaxilla immediately lateral to the middle of the rostrum, and as in
Nothosaurus tchernovi, the two anteriormost fangs fit between the two anteriormost symphyscal fangs when the jaws are closed." Nothosaurs flourished in
Europe during the m i d d l e Triassic, about 225 million years ago, and were
probably able to come out of the water, rather like modern pinnipeds do. T h e
nostrils, which would normally be at the tip of the snout, were actually set
further back, an adaptation that supports the suggestion that Nothosaurus was
at least semiaquatic. T h e y are classified as sauroptcrygians, a diversified group
that includes the placodonts and pachypleurosaurs and the later plesiosaurs.
Nothosaurus
was a swimmer,
but
it was capable oj
moving on land loo,
perhaps like some of
today's
tors.
water
Nothosaurs
preceded
and may
to them.
moni-
plesiosaurs
he ancestral
U s i n g a tail that
w a s twice as long as
its bead ana body,
1
Hovasaurus plied
the late
seas,
million
Adults
Penman
about
1^0
years ago.
were less
shows poor ossification of the l i m b girdles, suggesting that it could not walk
very well, but because its limbs were also not specialized into paddles, it
probably moved by undulations of the long tail. As O'Keefe (2002a) wrote,
"Plcsiosaurs were advanced over their 'nothosaur-gradc' forebcarers in the
evolution of wing-shaped fore and hind flippers that generated thrust via lift
as well as drag." Like other sauropterygians, the pachyplcurosaurs had an array
of gastralia, the belly ribs that formed a ventral support system under the
animal's underside and, along with the ribs, completely enclosed it in a shield
of internal bony armor. Carroll and Gaskill (1985) described Pachypleurosaurus
as having a moderately long neck and a particularly small skull, with widely
spaced, peglike teeth that suggested that it chased and caught fish. A tiny
nothosaur, possibly Pachypleurosaurus, was reported from Switzerland, with
large eyes and unossificd limbs that indicate that it was certainly an embryo
( S a n d e r 1988). Because it was not found in conjunction with an adult, this
suggests that the nothosaurs laid eggs perhaps on land.
T h e monotypical Pistosaurus is believed to be morphologically intermediate
between the nothosaurs and the plcsiosaurs proper. It is known only from the
With skeletal
characteristics
of
both
nothosaurs
and
plesiosaurs,
Pistosaurus may
have been
to
This
have been oceangoing creatures, but there has been no morphological analysis
creature lived in
of the way these animals moved, so we do not know whether it swam like a
crocodile (as nothosaurs d i d ) or paddled ( o r rowed) like a turtle or sea lion.
Placodonts were sturdy-bodied reptiles that fed on shellfish, harvesting
them with their shovcl-likc front teeth and crushing them between the heavy,
ancestral
the plesiosaurs.
middle
10-foot-long
the
Triassic,
about 1JO
years ago.
million
platelike teeth on the roofs of their mouths and their lower jaws. (Placodont
means "plate-teeth.") T h e i r feet were webbed and less flipper-like than those
of other marine reptiles, so they might have paddled in shallow water in
search of food, but like t o d a y s marine iguanas, they also might have spent
time out of the water. However, like the nothosaurs, their limb girdles were
too weak to support these large, heavy animals on land, so they may have
rested in the shallows when not feeding. Like that of the iguana, the tail of
Placodus was long and laterally compressed and probably helped in aquatic
propulsion. Placodus, which appeared and disappeared in the Triassic, was
characterized by a massive skull and the presence of massive gastralia. T h e r e
was a single row of dermal ossifications above the neural spines of the
vertebrae, which was probably expressed as a row of spines running down the
m i d d l e of the back. Placodonts looked something like overgrown, bucktoothed iguanas, and they came in armored and unarmored varieties. Placodus
may have reached a length of 6 feet, including the tail.
Henodus was a sort of armored placodont, with a body as wide as it was
long. At a length of about 3 'A feet, it had a boxlike head and a flexible tail,
which was probably used in locomotion, along with its feet. It looked something like a toothless Placodus pecking out from under a heavy, horny blanket.
Henodus was well armored, with protective shells top and bottom composed of
bony plates covered in horn. T h i s unusual marine reptile also had short foreand hind limbs, and its relatively small feet might have been webbed. Henodus
which means "one tooth" lacked the large, crushing teeth typical of the
placodonts, and it was originally assumed that its hard, horny beak was used
to get and crush shellfish, but recent studies have revealed numerous small
teeth that formed a combhkc structure that probably served as a sieving
apparatus for small crustaceans. Even though Benton (1990a) described it as
"startlingly similar in body form to the turtles," Henodus was not a turtle.
W h e r e a s the shell of a turtle develops largely from its skeletal bones most of
the carapace ( u p p e r s h e l l ) from the neural spines of the vertebrae and the ribs,
and the front end of the plastron (lower s h e l l ) from elements of the shoulder
girdle the a r m o r of Henodus developed exclusively from bony plates in the
skin,
ferent from and much more numerous than the plates of a turtles shell.
38
5 t A
WKAgoHS
The placodonts
("plate-teeth")
probably
used
their
Although it looks
something like a
turtle,
the j-foot-
long
Triassic
reptile
Henodus was
actually
an
placodont.
in
held a
sieve,
baleen-like
but how
Henodus
what
armored
Grooves
it fed on,Jor
mystery.
fedor
Germany.
In 1999, M i c h a e l Caldwell described a new species of a primitive marine
lizard known as Coniasaurus, from the chalk deposits at Brighton in southeastern England. ( A n earlier species, Coniasaurus crassidens, from the same
location had been described by R i c h a r d Owen in 1850.) T h e coniasaurs were
smallish lizards, probably no more than 3 feet long, with small heads and
elongated necks, bodies, and tails. A l t h o u g h they do not figure in the ancestry
of snakes (or m o s a s a u r s ) , the small size, elongated body, and small head
suggest a feeding strategy not unlike that of some extant sea snakes, that is,
probing in crevices in coral reefs and rocky shores.
W h e t h e r or not birds are dinosaurs, the ancient marine reptiles certainlv
were not. T h e y were not descended from dinosaur ancestors and represent a
completely different vertebrate lineage. In any case, the nonavian dinosaurs
were terrestrial and the marine reptiles were, well, marine. For an aquatic
existence, the marine reptiles had fins, while the terrestrial dinosaurs had legs.
Based on the structure of their pelvic girdles, dinosaurs have been divided into
two groups, the bird-hipped ( o r n i t h i s c h i a n ) and the lizard-hipped ( s a u rischian). T h e hips and hind limbs of dinosaurs are designed for weight
bearing and bipedal or quadrupedal upright walking, whereas those of the
ichthyosaurs and plesiosaurs could not possibly have borne any weight on
land. In fact, the hip bones of the ichthyosaurs are not connected with the
vertebral column at all; they arc only reduced vestiges, buried in the a b d o m i nal wall. ( M a n y living whales, which have no pelvic bones at all and no hind
legs, either have tiny, vestigial hind limbs buried in the muscle where the
hips would be.) T h e limbs of a plesiosaur demonstrate that it was a swimmer,
not a walker. Plesiosaurs had four limbs of approximately the same size and
propelled themselves through the water with some combination of flapping
and rowing motions. Ichthyosaurs were built not unlike dolphins, except that
the seagoing reptiles had hind limbs, and the dolphins have lost their hind
limbs altogether. Like dolphins, ichthyosaurs steered with their fins and
propelled themselves with their tail fins, but whereas the tails of d o l p h i n s and
whales are horizontal, those of the ichthyosaurs were vertical. To the best of
our knowledge, the dinosaurs were terrestrial except, of course, those that
returned to the water as birds (e.g., the hesperornithiformes). S o m e of the
nonavian dinosaurs may have ventured into the water, as do many present-day
terrestrial animals such as hippos, elephants, and penguins, but all the plesiosaurs and ichthyosaurs were aquatic and had flippers and other appropriate
skeletal modifications for the marine environment. Regardless of how well
adapted they seemed for a marine lifestyle, however, they all had to come to
the surface to breathe.
T h e nothosaurs, placodonts, ichthyosaurs, plesiosaurs, and mosasaurs
were oriented horizontally, like whales, dolphins, and most fishes, because
that is the best way to swim through the water. (A few species of living
fishes sea horses, for example orient themselves vertically, but they move
slowly and inefficiently, which is surely not what the great marine predators
d i d . ) T h e long-necked plesiosaurs almost certainly oriented their bodies
horizontally, with their long, flexible necks stretched out in front, swinging
in
plesiosaurs
is a l s o s t r o n g ,
1!
1101
vcl
ancestors remain unidentified, they were beginning to fill the same apex
predator positions in the water that the carnivorous dinosaurs did on land.
T h e ichthyosaurs lived from the early Triassic ( a b o u t 240 million years a g o )
until about 93 m i l l i o n years ago 32 million years before the mass extinctions
at the end of the Cretaceous period. During their tenure, they developed a
basic body plan that consisted of a dolphin-like shape; a tooth-filled beak; a
vertically oriented, lunate tail fin like that of a shark; and, in most of the later
species, a dorsal fin. T h e earliest ichthyosaurs were about a yard long, but they
later grew to impressive lengths, such as the 45-foot-long Shonisaurus, which
got as big as a h u m p b a c k whale, and the so-far u n n a m e d ichthyosaur from
British C o l u m b i a , which was even larger. T h e ichthyosaurs of the Jurassic
period are the most numerous, suggesting a flowering of these animals some
140 million years ago. T h e best-known genus was Ichthyosaurus, preserved
mostly in England, but also from southern Germany and western Canada.
Jurassic seas were also occupied by plesiosaurs, marine reptiles with long
necks, and their relatives the pliosaurs, short-necked animals with powerful
jaws and teeth. Giant marine crocodiles with long, eel-like tails competed for
prey in Jurassic seas, but unlike ichthyosaurs, which were pelagic and never left
the water ( a l t h o u g h they had to surface to breathe, like the dolphins they
resembled), the crocs occupied a nearshorc environment and probably came
ashore to rest and lay their eggs. T h e ichthyosaurs were gone for 25 million
years when the asteroid ( o r c o m e t ) hit in the area that would become the Gulf
of M e x i c o . T h e plesiosaurs were extinct by the end of the Cretaceous, the
same time that the last of the nonavian dinosaurs disappeared.
T h e dinosaurs, crocodiles, pterosaurs (flying reptiles), and marine reptiles
(ichthyosaurs, plesiosaurs, and m o s a s a u r s ) are all broadly classified as ruling
reptiles, but not all of them were dinosaurs. And although the dominance of
the sea by the great marine reptiles coincided with the dominance of the land
by the dinosaurs (and, to a lesser extent, the dominance of the air by pteros a u r s ) , the reptiles were as popular in the Victorian era as dinosaurs arc today.
T h e fashionableness of ichthyosaurs and mosasaurs was due almost entirely
to the fact that their discovery predated that of the first dinosaurs. T h e i r
remains were far more complete and therefore far more dramatic than the
first discoveries of fragmented dinosaur fossils. Because the marine reptiles
were the first "prehistoric" a n i m a l s uncovered in English fossil beds, they led
to a great upsurge in the study of extinct a n i m a l s . Public interest in the
ichthyosaurs, plesiosaurs, and mosasaurs declined after the nineteenth century, but recently there has been a revival of interest, probably sparked by the
discovery of various spectacular new species, such as gigantic mosasaurs from
Israel and N e w Zealand and ichthyosaurs that were larger than full-grown
sperm whales.
Just as there is an ongoing discussion about whether dinosaurs were w a r m blooded, there is a comparable question about the marine reptiles: how could
these "cold-blooded" reptiles possibly have lived and hunted in the heatsapping oceans if all their living relatives become slow and sluggish when the
ambient temperature drops? Because the ichthyosaurs, plesiosaurs, and mosasaurs were reptiles, it has been assumed that, like their living relatives, they
were cold-blooded. ( T h e blood of reptiles is not actually "cold"; some desertdwelling lizards have higher body temperatures than m a m m a l s of comparable
size.) M a m m a l s and birds are endothermic, which means that they produce their
own heat metabolically and maintain a high, constant b o d y temperature,
regardless of the surrounding conditions. If the temperature of most birds
and m a m m a l s falls a few degrees below normal, they die. M o d e r n lizards,
snakes, and crocodiles rely on the heat of the sun to w a r m up their bodies so
that they can perform their everyday activities. W h e n the temperature drops
too low at night in the desert, for example lizards become sluggish and
inactive. As might be expected, the larger the reptile, the longer it takes to
warm up the body, but the longer it can store heat within. W h e n paleontologist Robert Bakker applied this formula to some of the fast-moving d i n o saurs, he realized that, as reptiles, they could not rely on the sun to heat them
up because it would take too long, and besides, they would not function very
well on cool or cloudy days.*
* The original studies of the time it took large reptiles to warm up were performed in the
late 1 9 4 0 s by Charles Bogcrt and Edwin Colbert of the American Museum of Natural
History and by Raymond Cowlcs of the University of California. Working with alligators,
they saw that the smallest specimens could warm up quickly: a 7-inch-long specimen took
only 90 seconds to raise its body temperature iC, while it took a 30-pounder five times as
long. They wrote in 1 9 4 6 , "Continuing this line of reasoning, it would seem probable that in
cles, m a k i n g it possible for them to swim faster and more efficiently than their
colder-bodied relatives. S o m e fast-swimming fishes, such as tuna and billfish,
have similar arrangements, and they are among the fastest fishes in the ocean.
T h e l a m n i d sharks and these fast-swimming fishes have tails of a similar
design crescent shaped, with the upper lobe equal in size to the lower. T h e s e
similarities do not indicate a phylogenetic relationship between these sharks
and fishes, but they do suggest that in their history, the same systems evolved
convergently. W h a t do these fishes and sharks with efficient radiators and
homocercal ( e q u a l - l o b e d ) tails have in common? T h e y are powerful predators
and unusually fast swimmers. ( T h e m a k o shark and the sailfish are believed to
be the fastest swimmers of their respective g r o u p s . ) If we look at the body
plan of many of the later ichthyosaurs, we find the same deep-bodied shape,
tapering to a narrow tail stock and, in many cases, a homocercal tail fin.
Bony fishes, for the most part, are not warm-blooded. As M c G o w a n
(1991a) wrote, "Fishes, being surrounded by water, quickly lose the heat
generated by their actively contracting muscles. T h i s is because the heat that
leaves the muscles is lost when the blood passes through the gills. S o m e fishes,
like the tuna, are able to maintain some of their s w i m m i n g muscles at a
relatively high and constant temperature, through modifications in their
blood vascular system." But most fishes do not have heat exchangers, so how
do they function? "First," says M c G o w a n , "the costs of aquatic locomotion
are about one-tenth those of locomotion on land, one important factor being
that fishes do not have to spend energy in supporting their body weight.
Another consideration is that as s w i m m i n g speed increases, the flow of water
over the gills increases, facilitating the uptake of oxygen. A third factor that
needs consideration is the metabolic rate of high-performance fishes like the
tuna. . . . Given their elevated body rate, high body temperature, and high
activity level, the tuna should probably be regarded as being as endothermic as
a bird or mammal."
T h e ability to deliver live young underwater is another hint that many of
the marine reptiles were endothermic. No living reptiles lay eggs in the water
(reptile embryos must respirate through the egg's permeable shell), and although there are major differences between the living and extinct groups, this
is probably not one of them. T h e evidence that ichthyosaurs delivered live
inherited the fishlikc flexion of the body, walking compresses first one side of
the thorax and then the other, meaning that both lungs cannot be simultaneously inflated when the animal is walking, and should the animal break into
a run, breathing becomes impossible. Living amphibians cannot breathe and
run at the same time, a situation Cowen refers to as "Carrier's Constraint."*
But this constraint is not applicable to air-breathing marine reptiles; they
obviously do not have to breathe and "run" at the same time, because their
high-performance locomotion takes place underwater, and their breathing
occurs at the surface.
Cowen also addresses what he calls "the ichthyosaur problem," pointing
out that even though they were reptiles, their major propulsion came from the
tail, "anatomically rather decoupled from the main body by a narrow caudal
peduncle." Because ichthyosaurs are shaped remarkably like dolphins, it is easy
to suggest that they were fast, flexible swimmers like their m a m m a l i a n counterparts. "If the body was flexible," says Cowen, "ichthyosaurs really hadn't
solved Carrier's Constraint," because flexion of the thorax even if generated
from the "decoupled" tail would alternately close the lungs down and make
it difficult if not impossible for the animal to breathe. If they had no stamina,
but only ambushed their prey from short distances while holding their breath,
the streamlined shape would make ecological sense. However, Cowen prefers a
somewhat unexpected interpretation, based on observations of penguins and
dolphins, the closest analogues to the streamlined shape of some ichthyo-
* Mammals can breathe and run at the same time because their four legs support the body
equally and the thorax is not twisted, says Cowen. "In fact, quadrupedal locomotion
encourages breathing on the run. The backbone flexes and straightens up and down with
each stride, alternately expanding and compressing the rib cage evenly. So horses, dogs, and
jackrabbits running at full speed take one breath per stride." Marine mammals also avoid
the lateral thoracic compression because their undulations are dorsoventral, as might be
expected in animals that arc descended from terrestrial mammals. Contra Cowan's hypothesis, however, there arc several species of monitor lizards that arc, according to a letter from
Colin McHenry, "lizards who think they are mammals. . . . T h e pcrentie
(Varanusggantrus)
acts like its on speed. Check out the pcrentie pens in any reptile park [McHenry lives in
Australia] and they'll be racing around their pens non-stop, more like a dog than a lizard."
saurs. ( T h e marine reptiles were quite diverse, and not all of them would
behave in the same way.) He wrote:
Ichthyosaurs are not penguins or dolphins, but it is possible that leaping
could provide yet another way to avoid C a r r i e r s Constraint. An ichthyosaur could swim at high speed in an undulating path, with the pectoral
fins providing upward or downward forces as needed. W i t h appropriate
control, the ichthyosaur could "porpoise," lifting clear of the water, pointing straight ahead, and in that position, could breathe at high speed,
exactly as [penguins d o ] . Propulsion by the laterally flexing body would be
resumed during the underwater phase, with the body traversing the highdrag surface zone at a reasonably high angle. In this situation, the performance of an ichthyosaur could certainly approach that of a dolphin of the
same body size.
Cowcn, a senior lecturer in g e o l o g y at the University of California at Davis,
added limericks in the margins of History of Life, perhaps as m e m o r y aids for
his students. Here's the one about Carrier's Constraint:
Fast-swimming air-breathers are rare
But ichthyosaurs d i d it with flair
T h e y swam up in a leap
(It's energetically cheap)
And they took a deep breath in mid-air
Although "Carrier's Constraint" is applicable to terrestrial reptiles, it
probably had nothing to do with the ability of marine reptiles to s w i m
efficiently at high speed and still be able to breathe. Dolphins, penguins, and
sea lions, and probably ichthyosaurs and some plesiosaurs, are all subject to
"drag" (loss of m o m e n t u m ) as they move through the water. As M c H c n r y
wrote (personal communication 2002):
As soon as a swimming animal approaches the surface, the drag it incurs
increases significantly. If dolphins, for example, came up only to get their
blowholes out of the water (as they do when s w i m m i n g at a more leisurely
p a c e ) , then they'd end up spending a great deal of their time in the high
drag zone near the surface. To approach the surface in a controlled manner, exhale and then inhale without drowning, and then dive again, would
all take time. By porpoising, the animal bursts through the high-drag zone
at high speed, and then enters a zone of zero drag the atmosphere. Its
m o m e n t u m gives it plenty of time to breathe before it re-enters the water at
high speed, allowing it to quickly get down below the high-drag zone and
keep its speed up.
S o m e recently evolved marine reptiles had little trouble making the transition from land to water. Terrestrial snakes are ectotherms, of course, and
because surface area relative to weight is higher in long, thin animals than in
shorter, rounded ones, their serpentine bodies give them a great deal of
control of their heating and cooling rates. Because ectotherms rely on external
sources of heat, they do not have to expend energy in keeping their bodies
w a r m and therefore require much less food than comparably sized m a m m a l s
or birds do. ( S o m e living reptiles, such as large crocodiles or very large snakes,
m a y eat only three or four times a year.) Sea snakes, therefore, simply moved
into the water with their metabolic heritage intact, and because water holds
heat much better than air does, they were able to benefit from the relatively
stable temperatures of w a r m tropical seas. And like their terrestrial relatives,
sea snakes don't have to eat very often.
M o s t fossils consist of bones, shells, or teeth and rarely reveal the soft parts.
O u r knowledge of the muscles, ligaments, skin, viscera, eyes, and circulatory
systems of long-extinct animals usually consists of educated guesses based on
osteological analysis. We can tell how big an animal was, and whether it was a
tree-climber or a cursorial predator, but we cannot tell how its circulatory
system worked or what kind of noises it made. Available spaces can tell us a
lot, such as the size of the brain that could be contained in a particular skull,
or what sort of bone structure would support what sort of muscles, but
beyond that, we enter the realm of speculation. One subject rarely discussed
in relation to marine reptiles is fat (which, of course, would not fossilize), yet
it is extremely important in the lives of some marine m a m m a l s . Indeed, it is
The
heaviest
reptile,
back
living
the leatherturtle
(Dcrmochelys
coriacca)
reach a
jeet
and
can
length of 8'A
weigh
more
sea
not the metabolism of whales and d o l p h i n s that enables some of them to live
in cold water but rather the thick envelope of blubber that encases them and
as 3,000 feet.
keeps the body heat in and the ambient cold out. S p e r m whales, among the
deepest divers of all whales and therefore subjected to the most intense cold,
have a thick blubber layer that enables them to hunt at depths up to a mile.
Other cetaceans such as bowheads, narwhals, and belugas live their entire lives
in arctic ice-filled waters with no ill effects. S o m e pinnipeds, such as walruses
and elephant seals and even some of the larger penguins, have evolved an
insulating layer of fat that enables them to live in cold water, and even though
the Cretaceous so,is mav have been ,1 lot warmer than the waters oi modern
Greenland, it is possible that some of the larger ichthyosaurs, plcsiosaurs, and
mosasaurs had a layer of blubber. Besides insulating them, it would have
provided additional buoyancy as they floated at the surface to breathe.
T h e leathcrback, the largest and most widely distributed of all living sea
turtles, is well known for its frequent appearance in cold water. T h e FAO
species guide Sea Turtles of the World ( M a r q u e z 1990) lists as its high-latitude
water; the only energy that is needed is that required to propel the animal
through the water. On land, the reptile needs energy to overcome the pull
of gravity and to hold the body off the ground, and energy to move the
limbs during locomotion. Another possible clue as to why reptiles adapted
to life in the sea is the abundance of fish, squid, and bclcmnites (a s q u i d like c e p h a l o p o d ) parts found in the stomachs of many marine reptile
fossils. T h i s suggests that the M e s o z o i c seas were an enormous larder of
animal protein waiting to be tapped. Ocean production is greatest in
shallow coastal water, so it is not surprising that marine reptiles evolved in
the shallow seas of the M e s o z o i c .
W r i t i n g of the Western Interior Seaway, N i c h o l l s and Russell (1990) said,
"Studies of oxygen isotope ratios in belemnites suggests a temperature
range . . . of 1727C [628oF] in the seaway between latitudes of 7}N and
4oN. W h i l e a thermal gradient was present in the seaway, it was not as
pronounced as today and there is no evidence of cold, arctic conditions." If
the ancient seas were much warmer than thev are todav, the reptiles
would
have had no real problem maintaining their metabolic levels. Sea snakes,
which live only in the tropics, obviously do not have a problem with cold
water. In answer to a question I asked h i m about the metabolism of marine
reptiles, Carpenter wrote:
Throughout most of the M e s o z o i c , global temperatures were much higher
and nowhere is there evidence of continental glaciation. Early Triassic:
Coal found in Antarctica, situated pretty much where it is now, implying
warm, wet conditions. Evaporites (salt, g y p s u m , etc.) deposits formed
under arid conditions c o m m o n in western U S , Europe and M i d d l e East.
Late Triassic: Initial rifting of Pangea. Development of warm, shallow seas
between Europe and N.A. Giant amphibians in Antarctica. Early Jurassic:
Continued rifting. Abundant coal in Antarctica, Australia, Asia. Evaporites along North Africa. Late Jurassic: Continuation of above, large
Sahara-like deserts in Western U S . Oceanic anoxic events in deep sea
indicate high sea surface productivity, hence high sea surface temperatures.
Ferns occur as far as 6o north, farther than their present range. Cretaceous: Oxygen isotope data indicates sea surface temperatures of 8095F
" T h e gist of the data," said Carpenter, "is that marine reptiles had no problems as you raise." Even if the surface temperature of the water was as high as
95F, the depths w o u l d be considerably cooler, so the marine reptiles could
have dived in pursuit of prey, then returned to the surface to digest it and
reenergize their systems. M a n y of these animals were the size of whales, which
means that their body mass would have held a lot of internally generated heat.
H e a t exchangers would have enhanced their performance as in the case of
today's tunas and mackerel sharks which would mean that the ichthyosaurs,
plesiosaurs, and mosasaurs had been even more effective as predators. It is
unlikely that the explanation for the activity level of the marine reptiles will be
found in a single factor; rather, it was likely a combination of internal
metabolism, large size, warmer water, and increased activity.
W h e n I asked M i c h a e l Caldwell why he thought the marine reptiles could
adapt to the u n w e l c o m i n g conditions of cold water, he answered: enzymes. He
went on to write:
M o s t morphologists, paleontologists, etc. forget about biochemistry. Enz y m e activation levels (concentrations and temperatures) are the key to
successful m e t a b o l i s m in varied environments.
Enzymes are proteins. T h e y are produced directly by nucleotide
sequences and can therefore rapidly evolve ( u n l i k e morphology, which is
the product of major genetic and epigenetic evolution). Therefore, organisms which evolve new enzymes can readily adapt to specific problems, e.g.,
temperature or other metabolic needs like increased s w i m m i n g speed.
T h e r m o r e g u l a t i o n per se, is not the issue. An arctic cod can swim in
4 C water. In water that w o u l d inactivate your enzymes, the fish muscle
contracts and moves and works extremely well. T h e question is not the
success or failure of e n d o - versus ectothcrmy, but rather the variety of
enzyme systems and whether they work in cold temperatures or hot.
Ectotherms with such competent enzyme systems are actually much
better adapted to thermally stable ( h o m c o t h e r m i c ) environments than are
e n d o t h e r m s . H e a t is not an issue the way it is for m a m m a l s . To function successfully, aquatic e n d o t h e r m s must insulate and eat tremendous
amounts of food. In short, it is rather surprising that endotherms returned
reptiles had left the scene forever were descended from terrestrial forebears,
and all took the remarkable step of returning to the sea from whence they
came. In the process, they acquired ( o r reacquired) profound adaptive m o d i fications that enabled them to engage in a fully or partially aquatic lifestyle.
W h e r e there had been legs there were now flippers. In most marine reptiles,
scales were lost in favor of a slick skin that would pass smoothly through the
water. S o m e developed long, narrow, tooth-studded jaws, the better to gobble
up slippery fish and squid, while others followed a different path and developed huge heads and powerful jaws that were suited for killing and crushing
prey as large as or larger than themselves. Still others grew impossibly long
necks that somehow enabled them to snatch fish that were jo feet away. Vision
and hearing were modified for lire in a viscous m e d i u m that transmits light
poorly but sound wonderfully well. ( W e do not know if any of the marine
reptiles echolocated, but we do know that dolphins, the m o d e r n - d a y analogues of ichthyosaurs, have perfected a system of sound transmission and
reception that far surpasses anything that Homo technologies has managed to
come up w i t h . ) At least some of them renounced e g g - l a y i n g a plan that
most m o d e r n reptiles adhere to and, like the whales and dolphins, learned
to give birth to living young underwater.
T h e largest living marine reptiles are the saltwater crocodile and the leatherback turtle, descendants of ancient lines whose origins predated that of the
ichthyosaurs, plesiosaurs, and mosasaurs. Living birds are now considered the
lineal descendants of dinosaurs, so we can see "dinosaurs" just by looking at
our bird feeders. Like so m a n y large, d o m i n a n t groups of animals in the
history of life on Earth, the large marine reptiles are gone, and they left no
descendants. ( T h e i m a g i n a r y Loch Ness monster is not descended from a
plcsiosaur or anything else, for that matter.) T h e r e are no saucer-eyed, fourflippercd ichthyosaurs; there are no 50-foot lioplcurodons or kronosaurs (for
which we can probably be grateful); and there are no huge mosasaurs, crushing a m m o n i t e shells in their double-hinged jaws. T h e r e arc living monitor
lizards, of course, and although the Komodo dragon (Varanus kotnodoensis) is a
formidable beast, it is a pussycat compared with the lion that was Tylosattrtts.
(In Pleistocene Australia there lived an e n o r m o u s m o n i t o r lizard that was
twice the length and weight o f the K o m o d o dragon; Megalania prisca reached 22
feet long and weighed about 1,300 pounds. It became extinct 10,000 years ago.)
If snakes are closely related to the g r o u p including mosasaurs, they may be
heirs to a great tradition of aquatic reptiles that once dominated the seas, but
even the largest pythons and boas are but legless shadows of the m i g h t y
mosasaurs.
T h e apex predators of today's oceans are considerably less fearsome than
those that swam in M e s o z o i c seas. T h e ne plus ultra of today's marine predators
is the great white shark, usually an cater of fish and marine m a m m a l s but
whose reputation for anthropophagy has been greatly enhanced by the author
of Jaws. T h e killer whale, which hunts in packs and grows much larger and
heavier than the white shark, is a more efficient and dangerous hunter, but
only of selected victims, ranging from salmon and squid to seals, sea lions,
porpoises, and even the great whales. S p e r m whales, at 60 feet and 60 tons, are
the largest predators alive today, but they feed mostly on squid. T h e r e were no
humans around to see the great marine reptiles, and we know of their e x i s tence only from the fossil evidence. In our mind's eye, we can conjure up
images of the swift ichthyosaurs coursing after fishes and squid; plesiosaurs
stretching their necks out to g r a b at passing prey; monstrous pliosaurs with
their bone-crushing jaws, grabbing at anything and everything that came
within range; and 40-foot-long mosasaurs, lizards that menaced the life of the
inland seaways. Just as the terrestrial dinosaurs ruled the land and the flying
reptiles dominated the skies, the marine reptiles were the lords of the subaqueous realm. T h e sea dragons have been gone for 65 million years, but from
the fossils, some of them beautifully preserved, we can make them live again.
Enter their ancient underwater world and sec how they lived, how they
hunted, how they gave birth, how they died.
The Ichthyosaurs
If a casual observer looked at a drawing of an ichthyosaur and identified it as a
dolphin, the mistake could be easily forgiven. A closer examination would
reveal small hind flippers and, more critical, that the tail fin, instead of being
horizontal as it is in cetaceans, is vertical, as it is in sharks. T h o s e differences
notwithstanding, the ichthyosaurs of the m i d d l e Triassic through the Cretaceous demonstrate an incredible example of convergent evolution, with similar traits developing in totally unrelated groups of animals. In this case, the
ichthyosaurs would adapt to a life in the ocean, flourish, and die out 40
million years before the earliest cetaceans. In shape, they were streamlined like
From top: An
ichthyosaur,
porbeagle shark,
bottlenose
dolphin,
similar
evolutionary
solutions to the
problem
of moving
swiftly
through
the water.
The
ichthyosaur,
the
propulsion,
although
the
vertebral column
in
the ichthyosaur is in
the lower lobe of the
The
tuna
is probably the
best
designed of them
all
and
is generally
the fastest fishes, and they breathed through nostrils located close to the eyes,
not on the tip of the snout. U n l i k e the plesiosaurs, the ichthyosaurs d i d not
use their forclimbs for propulsion but used their lunate caudal fins, much the
way sharks do today. T h e y were reptiles and therefore had well-developed ribs
from their necks to their tails some of the post-Triassic ichthyosaurs had
more than 80 pairs of ribs which suggests a rigid trunk, with propulsion
deriving from undulations of the vertical tail fin.* (Early ichthyosaurs, such as
Chaobusaurus, had shorter ribs and probably swam by undulating the entire
body.)
T h e origin of ichthyosaurs is unknown, but their shaftlike limbs indicate
that they were certainly descended from terrestrial reptiles. C o l b e r t (1965)
wrote that "the ichthyosaurs comprise an isolated order of reptiles ultimately
derived from the cotylosaurs," which g r o u p Carroll (1988) describes as "a wide
range of primitive tetrapods . . . including a host of primitive amniotes," which
is not particularly helpful. " T h e typical ichthyosaurs of the Jurassic are so
highly modified for marine life" wrote R o m e r (1948), "that they show no clear
indications of relationship to any other reptilian group." T h e sudden a p pearance of the ichthyosaurs in the fossil record, with nothing even remotely
resembling a transitional form, has provided much support for the creationists'
dismissal of evolutionary theory. In Dinosaurs by Design, Duane T. Gish quotes
Edwin Colbert, Alfred Romer, and assorted other scientists, all of w h o m say
that there arc no clues to the ancestors of the ichthyosaurs. T h i s , says Gish,
shows that . . . explanations of the evolutionary process are really fairy
t a l e s . . . . If ordinary land reptiles changed into marine reptiles, then one of
these land reptiles must have ventured into the water, and after eons of
time and a long scries of genetic mistakes, it g r a d u a l l y changed into a fish* I he presence of multiple pairs of ribs is one of the defining characteristics of reptiles,
which have many more ribs than mammals do. Mammals have separate ribs only on the
vertebrae connected to the sternum - the so-called thoracic vertebrae. Reptiles have ribs on
the neck and tail vertebrae as well. In reptiles that have distinct necks, such as plesiosaurs
and dinosaurs, the neck ribs are often but not always short and reduced. In reptiles with
little or no distinct neck region, there arc long ribs on almost all vertebrae. This arrangement can be found in ichthyosaurs, mosasaurs. pliosaurs, and snakes.
like reptile or Ichthyosaurus. If this were true, we ought to find at least some
in-between kinds. Perhaps an animal with feet and legs gradually changing
into paddle like fins would give evidence of this evolution. But not one
such transitional form has ever been found! Every one of the marine
reptiles just popped into the fossil record fully-formed at the very s t a r t . . . .
T h i s is exactly what we w o u l d expect to find if God is the Creator of these
creatures. T h e s e fossils give powerful evidence against evolution.
Here, however, we will assume quite the opposite: these fossils constitute
powerful evidence for evolution. As with the m a m m a l s that evolved into
whales and dolphins, some of the reptiles returned to the sea and, in time,
acquired those characteristics that m a d e life in the water possible: a fishlike
shape, flippers instead of feet, a mouthful of sharp teeth for capturing slippery prey, and the ability to deliver their young alive in the water. Although
the evidence exists only for Stenopterygius, many of the ichthyosaurs are believed
to have had a dorsal fin to provide stability while swimming, since it has
developed independently in cetaceans, sharks, and bony fishes. Like the cetaceans that were to follow them and to which they are in no way related ichthyosaurs retained their air-breathing requirements and were therefore restricted in the a m o u n t of time they could spend underwater.
T h e ichthyosaurs lived from the early Triassic (about 250 million years
a g o ) and went extinct about 93 m i l l i o n years ago. During their 150 m i l l i o n year history, they developed many varied forms, all of them conforming to
the same basic body plan, with thematic variations. S o m e were small and
snub-nosed; others had long, pincer-like jaws; still others had an overhanging
upper jaw like that of a swordfish. T h e earliest ichthyosaurs were smallish, no
more than 3 feet long, but there were some monsters that reached a length of
50 feet, and some were even larger. We know that they were all aquatic
because, as M i c h a e l Taylor (1987c) wrote, "Paleontologists find ichthyosaur
fossils only in coastal or marine rocks." Besides, unless it was a snake, an
animal without functional legs w o u l d have had a rather difficult time on land.
In the M a r c h 6, 1999, issue of New Scientist, Kate Douglas wrote about
ichthyosaurs in an article called "Dinodolphin," which included this discussion of the adaptive process:
that time, and as the true nature of m a n y fossils was not generally understood,
L h w y d believed that the bones belonged to fish and called them Ichtbyospondyli."
W h i l e w a l k i n g on a Yorkshire beach in 1758, C a p t a i n C h a p m a n and M r .
W o o l e r came upon some fossil bones e m b e d d e d in a cliff. T h e y identified the
bones as b e l o n g i n g to an " a l l c g a t o r " and sent their written descriptions to the
Royal S o c i e t y of L o n d o n , where they were read into the Philosophical Transactions. ( T h e fossil, classified as the primitive crocodilian Steneosaurus bollensis,
is
C.H T.H
T/
0 5 A
U%.S
67
on the salamander Proteus, but the name Ichthyosaurus ("fish l i z a r d " ) had precedence and was subsequently applied to all the fossils. Five years later, in 1824,
the Reverend George Young found a fossil "fish lizard" at W h i t b y in Yorkshire. C a l l e d by Roger Osborne "a great figure in the beginnings of geology in
Yorkshire," Young described it as having the features of a crocodile, a fish, and
a dolphin, but "to what class of a n i m a l s this skeleton and others found at
W h i t b y should be assigned, it is difficult to determine." From the drawing in
his 1820 description (the actual fossil cannot be found), the animal has been
identified as Leptonectes acutirostris, the ichthyosaur now known as
Temnodon-
and a specimen sent to England. Home wrote, "f by no means consider it wholly a fish, but
rather view it in a similar light to those animals met with in New South Wales, which
appear to be so many deviations from ordinary structure, for the purpose of making
intermediate connecting links, to unite in the closest manner the classes of which the great
chain of animated beings is composed."
this. T h e forces ranged against him of custom, history, doctrine and c o m mon acceptance were just too formidable." T h e concept of extinction was
completely alien to people of Young's time and disposition.
M a r y Anning also discovered the first complete skeleton of the reptile
Conybeare and De la Beche n a m e d Plesiosaurus dolichodeirus. T h o m a s Birch
bought it from M a r y and m a d e it available to Conybeare, but it ultimately
ended up in the collection of the Duke of Buckingham. M a r y and Joseph
Anning made many important fossil finds, but it was M a r y who was famous
throughout Europe for her knowledge of the various fossilized animals she
was uncovering. As quoted in H u g h Torrens's 1995 biographical sketch of
M a r y Anning, Lady H a r r i e t Silvester wrote in her diary in 1824:
T h e extraordinary thing about this young woman is that she has m a d e
herself so thoroughly acquainted with the science that the m o m e n t she
finds any bones she knows to what tribe they belong. She fixes the bones on
a frame with cement and then makes drawings and has them engraved. . . .
It is certainly a wonderful instance of divine favourthat this poor, ignorant girl should be so blessed, for by reading an application she has arrived
at a degree of knowledge as to be in the habit of writing and talking to
professors and other clever men on the subject, and they all acknowledge
that she understands more of the science than anyone else in this k i n g d o m .
In addition to the ichthyosaurs and plesiosaurs, M a r y found Britain's first
pterodactyl. In 1838, for services rendered, M a r y Anning was given an annual
stipend from the British Association for the Advancement of Science, and she
was made the first honorary member of the Dorset C o u n t y M u s e u m in 1846,
a year before she died of breast cancer.
M a r y Anning is the reigning heroine of British paleontological history, but
her life story is somewhat more complicated than the one usually told to
schoolchildren. She was indeed the most famous fossil collector of her time,
but her story has been oversimplified and romanticized because it was an
uplifting Victorian tale of a young girl who discovered a "crocodile" in the
cliffs of Lyme R e g i s and sold it to an understanding gentleman just in time to
save her family from the poorhouse. According to Chris M c G o w a n (2001),
"Fossils were M a r y Anning's salvation, and not simply because they kept her
T 11 E
69
ne
basking reptile. Owen, the most celebrated anatomist of his time, originally
believed that the downward-pointing tail of the fossil was a p o s t m o r t e m
artifact and that it should have been straight, so early-nineteenth-century
illustrations of ichthyosaurs showed them out of the water with straight tails.
He later concluded that the tailbend existed in living ichthyosaurs and wrote,
" T h e appearance on the tail of the Ichthyosaurus . . . is too uniform and
common to be due entirely to an accidental and extrinsic cause. I am therefore
disposed to attribute it to an influence connected with some structure of the
recent animal; and most probably to the presence of a terminal caudal fin." It
was obvious that the downward tailbend was characteristic of the later ichthyosaurians and that it supported the lower lobe of the tail, unlike the caudal
vertebrae in sharks, which extend into the upper lobe of the tail.*
In 1846, Chaning Pearcc was examining a newly excavated fossil of Ichthyosaurus communis from Somersetshire when he found "a series of small
vertebrae lying on three or four posterior ribs; on removing another portion
* In .111 article devoted specifically to the tailbend in ichthyosaurs, Stephen J. Gould ( 1 9 9 0 )
documents the history of the discovery and recognizes the role that Owen played, but he
chooses to see the shape of the ichthyosaur as a convergence with fishes, as in "ichthyosaurs
are most celebrated for their convergence upon the external form of superior swimmers
among fishes." But if there is any convergence, it is surely with dolphins which developed
1 0 0 million years later. As Simon Conway Morris ( 2 0 0 2 ) wrote (of the similarity between
ichthyosaurs and dolphins), "the streamlined bodies of the mammal and the reptile cleave
the ocean in much the same way, even though the ichthyosaur evolved from something like a
lizard and the dolphin from something like a dog. It's a textbook example of convergence."
of the clay the rami of the jaws, and other parts of the head were visible." He
described how the tiny skeleton lay in the body cavity of the larger one and
observed, "while the posterior two thirds of the little animal is within the
pelvis, the head appears to protrude beyond it, and apparently being expelled
at the time of death." After consulting with R i c h a r d Owen, Pearce concluded
that he was indeed l o o k i n g at the frozen m o m e n t when an ichthyosaur
mother had died in the process of giving birth. In his 1880 " R e p o r t on the
M o d e of R e p r o d u c t i o n of Certain Species of Ichthyosaurus from the Lias of
England and Wurtemburg," H a r r y Govier Seeley, professor of geology at
King's College, L o n d o n ( a n d author of the first definitive work on flying
reptiles), examined the various fossils that appeared to contain miniature
ichthyosaurs. S o m e of these tiny skeletons associated with mature ones had
been interpreted as stomach contents, as if the larger ones had eaten the
smaller ones, but when skeletons were found in the pelvic region that is,
beyond the stomach in the digestive process it became clear that these were
fetuses and that ichthyosaurs gave birth to living young underwater, much the
way whales and dolphins do today. In England and Germany, Seeley examined as m a n y of the fossils-with-embryos as he could and wrote:
I therefore submit that the evidence indicates that these Ichthyosaurs were
viviparous, and were probably produced of different relative bulk in different species; and it m a y be from feeble health of the parent or from some
accident of position in the young that they were not produced alive, and
thus have left a record of their m o d e of reproduction to which no allied
extinct g r o u p of animals has shown a parallel. T h e r e is some evidence that
in certain cases many young were produced at a birth, and although the
specimens are not in the best state of preservation, a n a l o g y strongly suggests that this is a distinctive character of certain species.
Ichthyosaurus communis (the "common ichthyosaur," if you w i l l ) is the species
m o s t often found in the limestone of L y m e R e g i s and Street. It was a
m e d i u m - s i z e d ichthyosaur with a sharply pointed snout; a tuna-like, lunate
tail fin; and probably a prominent dorsal fin. In a specimen found at Kilve
( S o m e r s e t ) in 1985, a tiny embryo was found in the immediate vicinity of a
fossilized adult; the proportionally large skull, head, and orbit of the little
one m a d e it clear that it was an embryo. (In a 1993 discussion, Charles
Deeming, Beverly Halstead, M a n k a t o M a n a b e , and David U n w i n described
"the reproductive b i o l o g y of ichthyosaurs" quite an extraordinary a c c o m plishment with regard to a n i m a l s that have been extinct for 85 million years.)
Because cetaceans usually deliver their young tail first, so that the neonate
remains attached to the female as long as possible and will not drown if there
is a hitch in the birthing process, the same assumption was applied to ichthyosaurs. But because many of the "small associated" individuals are preserved in the headfirst position, D e e m i n g et al. assumed that in these cases,
"complications during the birth process (perhaps the embryo was too large
The stripes on
for the pelvic opening and lodged in the birth c a n a l ) led to the death of the
this
Ichthyosaurus
corpse would almost certainly have caused the death of the mother." T h i s
mother
conjectural,
mother in the process of giving birth; a n o r m a l birth would not have killed
the mother, and there would have been no reason for the fossilization of the
pair, but if a difficult birth resulted in the death of the mother, the fossils
hut it is
to
under-
oj about
But for all the "knowledge and cleverness," lchthyosaurology in H o l z maden was in a chaotic state. As Axel Hungerbiihler wrote in a 1994 study,
"when the Lower Jurassic ichthyosaurs of G e r m a n y were reviewed by M c Gowan, he found it impossible to identify most of the specimens described
by [German geologist Friedrich] Quenstedt. . . . Labels on most of the type
specimens* were missing and the type material is scattered throughout the
collection. But above all, it is Quenstedt's idiosyncratic approach to taxonomy
that has caused the most taxonomic confusion for later authors." Quenstedt's
descriptions, often resorting to t r i n o m i a l s and even q u a d r i n o m i a l s , were
published in the m i d to late nineteenth century, and when M c G o w a n examined the material in the m u s e u m at Tubingen, he wrote that of the 28 names
erected, only ten were valid, and to these ten he added two new ones, Stenopterygius cuniceps and S. macrophasma. T h e osteological and taxonomic reasons for
McGowan's revisions are too technical for this discussion, but when H u n g e r biihler examined the specimens whose names and identifications Quenstedt
had mangled, he wrote that "the taxonomy of Toarcian ichthyosaurs is far
from certain and further investigations arc needed to establish the validity of
a number of species from Germany."
Christopher M c G o w a n of the Royal O n t a r i o M u s e u m in Toronto has
devoted his professional life to the study of ichthyosaurs. Born and educated
in England, he wrote his doctoral dissertation at the University of L o n d o n on
" T h e Cranial M o r p h o l o g y and Interrelationships of Lower Liassic Ichthyosaurs." Since then, he has published more than 40 papers on these marine
reptiles. In Dinosaurs, Spitfires, and Sea Dragons, which includes a comprehensive,
eminently readable study of the ichthyosaurs, M c G o w a n defined them chronologically, that is, ichthyosaurs of the Triassic (the earliest known, from 250 to
* Under the formal rules for naming species in the International Code of Zoological
Nomenclature, each species of animal or plant must be represented by a type specimen, or
holotype, which is the single specimen selected by the original describer of a species to be
the standard-bearer for the new name. In simple terms, it is the original specimen of a
species. The holotype specimen may or may not be the first ever collected, and it may or
may not be a good example of its kind, but it has the official designation. Thus, the first
time Conybeare decided to call a particular fossil Ichthyosaurus communis, that fossil became
the type specimen. These rules apply to recent as well as fossil species.
The
early
ichthyosaurs,
developed the vertical, sharklike tail fin a n d had a trailing tail, perhaps with an
incipient upper lobe. Like m a n y of the ichthyosaurs that followed, Mixosaurus
had e n o r m o u s eyes, set in a circular framework of bony plates known as the
sclerotic ring. Primitive crocodilians had sclerotic rings, but m o d e r n crocs and
the
such
Mixosaurus, did
not have the
pronounced
upper
alligators do not. Living fishes, birds, and some lizards have sclerotic rings, and
ones.
their function is to support the eyeballs. Birds and most reptiles do not have
muscles attached to their lenses and therefore cannot change the focus of their
found in
eyes. Instead, they compress the eyeball using the sclerotic ring, thus pushing
Indonesia,
the lens farther from the back of the eyeball. T h e principle is the same as using
North
the focus knob on a pair of binoculars to shorten or lengthen the focal length.
T h e term accommodation in this context refers to any m e c h a n i s m that allows
an eye to bring closer objects into focus on the retina. A n i m a l s generally
as
]-joot-long
Fossils oj this
China,
Europe,
America,
Spitsbergen.
and
The
yjoot-long
ichthycsaur
Triassic
Utatsusaurus did
not have the vertical
t a i l J i n that
characterized
later
species; it moved by
flexing
body.
its entire
It was
discovered
in
Japan.
causing it to change shape as well as moving the lens within it. (Even humans
sometimes squint in an attempt to correct poor focus, which also changes the
shape of the eyeball.) Conversely, some birds cormorants, for example
drastically change the shape of their lenses when they go underwater so that
they can compensate for the fact that their corneas no longer provide much
focusing power.* Because we cannot do this, we can't see well underwater
w i t h o u t trapping a layer of air around our eyes in a diving mask.
Like C h r i s M c G o w a n , Ryosuke M o t a n i , formerly of the Royal Ontario
M u s e u m but now at the University of Oregon, is a paleontologist who has
devoted h i m s e l f to the study of ichthyosaurs. ( H i s 1997 doctoral dissertation
at the University of Toronto was "Phylogeny of the Ichthyosauria with Special
Reference to Triassic Forms.") In their 1998 discussion of Utatsusaurus, M o t a n i ,
M i n o u r a , and A n d o wrote that the 5-foot-long ichthyosaur (which they called
"phylogenetically basal a m o n g ichthyosaurs") had about 40 vertebrae in the
front part of its b o d y the same n u m b e r as living catsharks ( S c i l i o r h i n i d a e ) ,
* M . P. Rowe ( 2 0 0 0 ) wrote that the conventional interpretation of sclerotic rings was that
they altered the focus of the eye by changing the shape of the eyeball. In some fishes and
early tetrapods, sclerotic rings occur along with double cones on the retina, indicating
enhanced color vision. We see such adaptations in birds, for example, and other animals
that are active during daylight hours. This suggests that ichthyosaurs and other marine
reptiles with sclerotic rings were active during the day and had some color vision. The large
eyes of ichthyosaurs suggest low-light visual acuity, especially helpful in the fast pursuit of
prey ( M o t a n i 2 0 0 2 ) .
which lack a high, vertical tail fin and move by flexing their entire bodies. T h e
eel-like s w i m m i n g style of the earliest ichthyosaurs suggests that they lived
and hunted in shallow water, where maneuverability w o u l d be more advantageous than in the open ocean, where greater speed is required. Later ichthyosaurs would develop a lunate, vertical tail fin like that of the speedy
mackerel sharks (great white, mako, porbeagle) and the tunas, which provides
propulsion with little or no body flexion. L i n g h a m - S o l i a r (1991a) identified
this type of locomotion as "axial oscillation" and wrote, "in this m o d e the
entire body, which is spindle or torpedo shaped, remains stiff or nearly s o . . . .
T h e force is transmitted from the massive musculature to the stiff caudal fin,
via a strong series of tendons."' T h e vertebrae of the early ichthyosaurs were
>
long and narrow, conducive to flexible, oscillatory swimming, but they became shorter and flatter in later forms, modifications that were required for
the swimming style of heavier-bodied, tail-propelled swimmers.
T h e ichthyosaurs for which fossils have been found were a large and varied
group, but they all conformed to a basic body plan (known to paleontologists
as a bauplan, from the German for "work p l a n " ) : streamlined body, long snout,
four flippers, and, in the later species, vertical tail fin and dorsal fin. T h e r e was
enough variation, however, to create many genera, which differed in such
particulars as length of rostrum, size of eyes, number of bones in the flippers,
and overall size. T h e first ichthyosaurs were long and skinny, almost eel-like in
form, but later species were deeper-bodied like the mackerel sharks, particularly the porbeagle (Laiima nasus) and the great white {Carcbaroion carcharias). "Sharks evolved several body forms," wrote M o t a n i et al. (1996),
"some of which are also found in ichthyosaurs. Because of these similarities,
sharks provide the best analogue for ichthyosaurs in overall body shape and
locomotion, although differing in details."
T h e earliest ichthyosaurs in contrast to the earliest discovered ichthyo* This description is equally applicable to some of the small, swift cetaceans, such as the
spinner, spotter, common, and boitlenose dolphins, all of which are as streamlined as
anything that swims and as fast as any fish in the ocean except perhaps the blucfin tuna.
The difference between cetaceans and ichthyosaurs, of course, is that the cetaceans move
their horizontal tails up and down, while the ichthyosaurs (and the sharks and fishes) move
their vertical tail fins from side to side.
saur fossils were small, with a five-fingered forcfin, a relatively short snout,
and no sign of the downturned vertebral column that would characterize later
forms. A m o n g these early Triassic forms are Grippia from Spitsbergen, Utatsusaurus from Japan, and Chaohusaurus from C h i n a . According to Massare and
Callaway (1990), "Triassic ichthyosaurs had less compact, more elongated
bodies than the Jurassic species, a factor that must have affected their swimming capabilities." Indeed, the long, unbent vertebral column probably meant
that they had not developed the bilobed caudal fin, probably swam with an
undulating motion like crocodilians or mosasaurs, and were probably ambush
predators. ( T h e alternative form of predation is "pursuit," whereby powerful
swimmers like the later ichthyosaurs and the pliosaurs actively chase their
prey over greater distances.) Massare and Callaway conclude their article by
noting, "the Triassic probably represents a period of experimentation and
fine-tuning for the ichthyosaurs."
"Primitive Early Triassic ichthyosaurs are rare," wrote M a z i n et al. in 1991;
"up to now only seven species, referred to six genera have been described."
T h e y are
Grippia (Spitsbergen),
Chaohusaurus ( C h i n a ) ,
Utatsusaurus ( J a p a n ) ,
Svalhardosaurus (Spitsbergen), Omphalosaurus ( N e v a d a ) , and two species of Chensaurus ( C h i n a ) . (Chensaurus is now considered a junior synonym for Chaohusaurus.) In 1988, three specimens of a new Triassic ichthyosaur were found in
T h a i l a n d , significantly expanding the database for the study of early ichthyosaurs. It had a long, pointed snout and high, conical teeth. T h e skull,
which was incomplete, was approximately 6 inches long, suggesting an overall
body length of 2 feet. T h e new species was named Thaisaurus chonglakmanii, for
T h a i l a n d and C h o n g p a n C h o n g l a k m a n i , the man who found the specimens
in a limestone q u a r r y near the city of Phattalung. But in his 1999 "Phylogcny
of the Ichthyopterygia," M o t a n i downgraded Thaisaurus to the ignominious
classification of incertae sedis ("affinities uncertain"), because it appears to be
very similar to Chaohusaurus geishanensis, and also because the material analyzed
by M a z i n et al. is "too poorly known to be included in the cladistic analysis."
Describing the discovery of a new specimen of Utatsusaurus in Japan in a
Scientific American article, M o t a n i (2000b) wrote: " W h e n I saw the skeleton for
the first time, I knew that Utatsusaurus was exactly what paleontologists had
been expecting to find for years: an ichthyosaur that looked like a lizard with
" There arc two regions of Germany where the close-grained stone preserves incredible
detail in fossils: Holzmaden, located about 20 miles southeast of Stuttgart, and the more
famous Solnhofcn, the home of Arebaeopteryx and Compsogmtbus, which is in Bavaria, about 6 0
miles northwest of Munich.
T h e identification, classification, and naming of ichthyosaurs is an ongoing enterprise; as various paleontologists examine and reexamine the m a t e rialwhich is often fragmentary and sometimes goes missing the phylogeny
of these marine reptiles is stretched, compressed, modified, altered, and corrected to such a degree that the subject can only be described as a work in
} lolzmaden
Shales
of Germany,
we
know that
Stenopterygius
had a large, lunate
tail and a large
dorsalfin.
Any attempt to produce a stable taxonomy is circumscribed by the uncertainty of recognizing natural groupings of individuals. It is impossible to
estimate the range of individual variations, and the effects of allomctric
It
was
dolphin.
Caypullisaurus?)
Georg Baur, a Ph.D. from Germany, worked for O. C. M a r s h in N e w
Haven, Connecticut, for many years and provided a much-needed backg r o u n d in b i o l o g y to Marsh's expertise in g e o l o g y and mineralogy. Like all
Marsh's assistants, Baur was underpaid, underappreciated, and restricted in
what he was allowed to publish under his own name. Still, in 1887, he wrote
"On the M o r p h o l o g y and O r i g i n of the Ichthyopterygia" in which he opined
that "the Ichthyopterygia were developed from land-living reptiles which very
much approached the S p h e n o d o n t i d a e . . . . T h e i r fins are not original but
secondary formations, like the paddles of cetaceans." T h e family S p h e n o d o n tidae is a mostly extinct g r o u p of land reptiles represented today by the
tuataras (Sphenodon), the only surviving members of the order Sphcnodontia.
A l l other m e m b e r s of the order ( a n d f a m i l y ) are known only from the
* T h e International Code of Zoological Nomenclature recommends the use of " M a c "
rather than " M c " in scientific names, hence the apparent misspelling.
Mesozoic. Sphenodon and the crocodilians resemble each other and differ from
all other living reptiles in that they have diapsid skulls, as did the ichthyosaurs. It was on the basis of the skull that Baur affiliated the ichthyosaurs
with the Sphenodontidae ("turning now to the upper part of the skull," he
wrote, "we find the parietal bones of exactly the same structure as in Sphenodon,
and in front of those very small frontals.") But M a i s c h believes that Baur ( a n d
others who suggest that ichthyosaurs are descended from d i a p s i d s ) was dead
wrong and that "the ichthyosaurian ancestor was an anapsid amniote or
proto-amniote."* Diapsid, anapsid, amniote, or proto-amniote, the ancestors
of the ichthyosaurs remain well hidden in the rocks.
T h e family Mixosauridae was erected in 1887 by Baur, and he included the
species that had previously been classified as Ichthyosaurus cornalianus and /.
atoms. T h e animal now known as Mixosaurus cornalianus was described from
specimens found in Basano, Italy, but they were destroyed when the M i l a n
M u s e u m of Natural H i s t o r y was b o m b e d during W o r l d W i r II. T h e best
material now available is from T i c i n o , Switzerland, and is housed in the
museum in Zurich, but as Callaway (1997b) points out, "this material remains
undescribed to the present date . . . because access to researchers has been
denied or severely limited." Nevertheless, C a l l a w a y managed to examine some
of the Zurich material and wrote "an interim update on various aspects of
Mixosaurus."
natans,
M.
nordeskioeldii,
and the various Phalarodon species into two recognized species, Mixosaurus
* Anapsids arc tctrapods characterized by the lack of temporal fencstrac, large holes in the
side of the skull. Whereas anapsids have no holes behind the eye socket, diapsids have two
on each side of the skull. The function of these holes has long been debated, but no
consensus has been reached. Many believe that they allow muscles to expand and lengthen,
which would result in a stronger jaw musculature, and the longer muscle fibers would allow
an increase in the gape. The taxon Anapsida includes turtles and their extinct relatives.
Amniotes are vertebrates that possess an extraembryonic layer called an amnion within the
egg or womb, which replaces the aquatic environment required for developing vertebrate
embryos. Amniotes may reproduce on land and may respirate without the assistance of a
body of water. Their development was a monumental event in vertebrate evolution, allowing for domination of the land and exploitation of the food resources growing there. All
birds, reptiles, and mammals are amniotes.
cornalianus and M. atavus. T h e mixosaurs have well-developed belly ribs ( g a s tralia), but what function they performed is uncertain. S o m e plesiosaurs also
had these tightly knit structures that formed a sort of plastron or bony shield
on the underbelly, but, Callaway writes, "they may simply represent a feature
retained from terrestrial ancestors, although they seem very well developed
for mere vestigial structures." He concludes, "Mixosaurus became the quintessential Triassic ichthyosaur used to illustrate both scientific and popular
literature. . . . M i x o s a u r s are not the most primitive of ichthyosaurs, as often
depicted in much of the older literature, but are in some respects as derived as
or more derived than Late Triassic and post-Triassic forms."
In 1998, M i c h a e l M a i s c h , a German ichthyosaurologist working in Tubingen, wrote that there might be another species of mixosaur from M o n t e
San Giorgio. W i t h Andreas M a t z k e , he created the new genus Wimanius, "for
Prof. Dr. Carl W i m a n of the University of U p p s a l a ( S w e d e n ) for his excellent
contributions to palcoherpetology, particularly on the Triassic ichthyosaurs."* M a i s c h also reexamined the specimen from H o l z m a d e n originally
designated Leptopterygius disinteger by von Huene in 1926 and renamed it Suevoleviatban from Suevo, Latin for S w a b i a in Germany, and leviathan, the Hebrew
name for "a sea dragon of the antediluvian earth." Known from only a single
specimen, the Swabian leviathan was a largish ichthyosaur, about 13 feet long,
with m e d i u m - s i z e d eyes. In their 2001 study, M a i s c h and M a t z k e recognized
Phalarodon ( N i c h o l l s et al. 1999) and wrote, "Here we focus on a third valid
ichthyosaur taxon, hitherto known as Mixosaurus major and usually dismissed
as a nomen d u b i u m (e.g. Callaway and Massare 1989). It is demonstrated that
* But when Motani ( 1 9 9 9 a ) reexamined the bone that was described as having teeth, he
noted that "this bone is possibly a broken pterygoid. Maisch and Matzke noted that the
only other ichthyoptcrygian with teeth on the palate was Crippia, and that the teeth were on
the palatine in that genus, citing Wiman ( 1 9 3 3 ) . They accordingly identified the bone in
their new genus as the palatine, however what was described as the palatal teeth by Wiman
have been shown to be the second row of maxillary teeth. The only ichthyopterygian with
teeth on the palate is Utatsusaurus, and these teeth are on the pterygoid. It is therefore
possible that the bone is actually a pterygoid rather than palatine." Although Maisch and
Matzke point out the differences between Wimanius and Mikadocepbalus, they share many
characteristics in common and may in fact be the same species.
the type material of this species, despite its fragmentary nature, is diagnostic
on the generic and specific level and can be referred to the genus Phalarodon M e r r i a m 1910, which was up to now u n k n o w n from the Germanic
Muschclkalk."
In 1998, Maisch and M a t z k e published the first description of "a crested
predatory mixosaurid from the M i d d l e Triassic of the Germanic Basin,"
which they identified as a new genus and called Conteclopalatus ("closed pala t e " ) . It reached a length of 16 feet, more than twice that of other mixosaurids,
but it differed from all others in the shape of its skull, which M a i s c h and
M a t z k e (2000) called "the most bizarre of any known ichthyosaur." It had a
high sagittal crest, which the authors interpreted as "correlated with a unique
arrangement of the jaw adductor muscles . . . with the internal jaw adductors
extending over most of the skull roof up to the external narial opening." T h e y
believed that this arrangement greatly increased the biting force of the jaws
and made Contectopalatus a particularly effective marine predator. It was long
thought that the mixosaurs were primitive versions of what was to follow, but
Contectopalatus was extremely specialized, capable of crunching the hard shells
of ammonites and perhaps even preying on smaller ichthyosaurs.
A (possible) later development in ichthyosaurs was durophagy, which
means the eating of hard objects, such as bivalves and ammonites. T h e teeth
of Omphalosaurus (omphalos is "navel" in Greek) consisted of an irregular pavement of button-like teeth set in short, massive jawbones. T h e original specimen was discovered in Nevada by J. C. M e r r i a m (1906), who believed that it
represented a distinct g r o u p of reptiles related to placodonts or rhynchosaurs.
In 1910, fossils were found in Spitsbergen that Carl W i m a n believed were the
limb bones of ichthyosaurs, and he assigned them to the species Omphalosaurus
nevadanus. McGowan (1991a) wrote that it was a "problematic and poorlyknown Triassic ichthyosaur," and according to Sander (2000), there is a
question of whether Omphalosaurus is really an ichthyosaur and whether the
limb bones of the Spitsbergen specimen actually came from the same animal
as the jaws. M o t a n i (2000a) argues that "there is insufficient reason to consider Omphalosaurus as ichthyoptcrygian . . . [because] the characters used to
unite the two groups are all inconclusive because none are unique to the two
and most are lacking in basal ichthyoptcrygians." (In i860, R i c h a r d Owen
1 H L
C.H TH r O $ AUK S
1
87
(named for
Shasta
County,
California)
probably
looked
which
imaginary.
shown
is
It is
chasing
squid.
arbitrarily changed Blainville's Ichthyosauria to Ichthyopterygia ["fish flipp e r s " ] , "a name which is often, though incorrectly, used to designate this
order of reptiles" [ C a l l a w a y 1997a].)
At the turn of the nineteenth century, paleontologist John C. M e r r i a m of
the University of California at Berkeley was working in northern California
and Nevada, ably assisted by Annie Alexander, a wealthy amateur collector.
Alexander sponsored many of M c r r i a m ' s expeditions, and in recognition of
her considerable contributions to paleontology and to her founding of the
M u s e u m of Paleontology at Berkeley he named the species Shastasaurus alexandrae for her in 1902 (Shastasaurus was named for Shasta County, California).
But even now, wrote M c G o w a n (1994b), with new material being uncovered,
Shastasaurus is not a well-understood genus. " M a n y of the species," he noted,
"were erected on inadequate material especially vertebrae and rib fragmentsand should be considered nomina dubia ('doubtful names'). A l though there is information for the skull, fins, and girdles, most of the
material is incomplete. T h u s there is no complete skull, forefin or hindfin,
nor is the vertebral column known much beyond the pelvic girdle." Beginning
in 1986, M c G o w a n led expeditions to W i l l i s t o n Lake, British Columbia, and
found "the most complete skeleton to date," which he named Shastasaurus
] CJ-t T.H
0 5 A
U%.S
89
Shonisaurus from the Nevada desert, and was by far the largest marine reptile
ever found. T h r o u g h the combined efforts of the Royal Tyrrell M u s e u m , the
N a t i o n a l Science M u s e u m of Tokyo, and the Discovery Channel, a team led
by N i c h o l l s managed to excavate the skull in pieces in 1999. T h e skull alone
was 18 feet long, and the largest piece (the braincase and o r b i t ) weighed 8,860
pounds. It was airlifted from the site by a cargo helicopter usually used to
carry trucks and other heavy equipment. N i c h o l l s has spent the last s i x years
painstakingly excavating and studying the ichthyosaur, overcoming countless
obstacles to extract the fossil from its limestone bed on a remote riverbank
flooded for part of the year. For her efforts, Nicholls was named a Rolex
laureate in 2000, which includes a stipend of $100,000, meaning that the Swiss
watch company is financing a substantial portion of her very expensive
excavations.*
W h e n N i c h o l l s publishes the results of her excavations, the specimen will
be recognized as the largest ichthyosaur, and probably the largest marine
reptile, that ever lived. But for now, the largest of the ichthyosaurs is Shonisaurus. N a m e d for the Shoshone M o u n t a i n R a n g e , where the fossilized remains of 40 ichthyosaurs were discovered in 1928 by S i m e o n M u l l e r in a
naturally eroded area of what is now Berlin-Ichthyosaur State Park, Shonisaurus
is the state fossil of Nevada. Excavations began in 1954 under the direction of
Charles C a m p and S a m u e l W e l l e s of the University of California at Berkeley.
At a length of 50 feet, this gigantic reptile was approximately equal in size to
an adult h u m p b a c k whale. Its long backbone bent downward at the posterior
end, supporting the lower lobe of a sharklike tail. Its jaws were greatly
elongated, with teeth only at the front, and its flippers were 5 feet long and of
equal size in the front and back. W h e n C a m p was excavating the specimens,
In December 2 0 0 1 , in response to my question about how the excavation was going,
Nicholls wrote to me: "Work on our giant ichthyosaur is coming along w e l l . . . . The skull is
almost finished. It is not as complete as we had hoped, as the front of it was so badly
crushed. But we have a good part of the back of the skull, and the palate is in good shape.
Sounds funny doesn't it, to be enthused about a 'good palate'? The tail is prepared, and we
are working on front limbs and ribs. Hope to have a preliminary publication on it submitted next year, and will tell you more about it when that is o u t . . . . I am not one to release my
data to the popular press before it is published."
This
unlikely
looking
creature
is
Shonisaurus. At
a known length of
he noticed that they were all aligned in more or less the same direction,
leading him to suggest that they might have been mass stranded, much the
way certain whales do today.
one of the
largest
marine
for the sudden mass mortality of the shonisaurs of Nevada. Stranding re-
reptiles
that ever
quires a shallow, coastal location, but examination of the site led Orndorff et
lived.
al. (2001) to conclude that "the ichthyosaur bones were deposited on a deep
unusual in that Us
ocean shelf environment." If they didn't strand, then how to explain the
enormous
It was also
front
Shonisaurus, but ill health prevented him from seeing it through to publication.
Just before his death in 1975, C a m p entrusted his manuscript and accompanying illustrations to his friend Joseph Gregory, which resulted in two publications in 1976 and 1980 (both appear under Camp's n a m e ) . In the 1980 publication, there was a reconstruction of the skeleton of Shonisaurus popularis,
depicted as a deep-bodied creature with huge fins and a skull that was nearly
as long as its downturned tail. T h i s image of the gigantic ichthyosaur appeared in numerous popular and scientific publications, and it was generally
acknowledged that Shonisaurus, with its narrow, pointed head and unnaturally
deep rib cage, was one weird-looking animal. In 1990, Bradley Kosch of the
Berlin-Ichthyosaur State Park in Nevada revised the interpretation of Camp's
skeleton, noting that "Camp's often reproduced skeletal reconstruction contains significant discrepancies from both his published description and his
unpublished field notes." Kosch realized that "as much as eight feet of the
dorsal region might not have been represented," and the ribs were too long,
which gave his restoration i t s exaggerated potbellied appearance. T h e n M c Gowan and M o t a n i (1999) remeasured the actual specimens housed at the
N a t u r a l H i s t o r y M u s e u m of the University of Nevada at Las Vegas and also
at the Berlin-Ichthyosaur State Park. C a m p had identified three species: Shonisaurus popularis (the most common, represented by 37 of the 40 individuals),
5. silberlingi, and S. mulleri. But after a careful examination of the fossils, M c Gowan and M o t a n i declared that "the likelihood of their having been three
species of Shonisaurus, the largest of the ichthyosaurs, seems unlikely," and there
was probably only a single species, 5. popularis.
W h e n M a r t i n S a n d e r of the Institute for Paleontology in Bonn examined
a fossil that had been exposed in the M u s c h c l k a l k beds of Karlstadt, he found
that it was a small shastasaurid, to date the most complete ichthyosaur
skeleton from that region of Germany. In 1997, he named it Shastasaurus neubigi,
for Bernd Neubig, who had discovered the fossil during railroad construction
work in 1985. In the early Triassic period, the Tethys Sea covered much of
what is now central northern Europe, and because large ichthyosaurs are
otherwise rare in the M u s c h c l k a l k Sea, S a n d e r opined that the individual
m i g h t have accidentally entered the shallow basin. Sander erected the species
Shastasaurus neubigi, "based on the structure of the vertebrae, in particular the
nature of the rib articulations, and of the pubis." Again, this illustrates the
ever-changing nature of paleontology, with some of the older specimens
relegated to the scrap heap of nomina dubia, some combined with others (the
correct term is synonymized), and others rejected completely because the original material was too fragmentary for an accurate diagnosis or simply because
a later worker decided that the earlier description was inaccurate. Have we
arrived at a point where we can say that we really understand ichthyosaur
phylogeny? As long as paleontologists keep digging, reexamining material in
existing collections, or arguing about which name ought to have priority, the
book on the fish lizards will remain open.
T h e shastasaurs some of which grew to enormous sizes represent an
example of convergent or parallel evolution with the toothed whales of the
Cenozoic era, although in terms of the rapid evolutionary tendency to hugeness, there is a similarity with the baleen whales, which likewise grew from
medium-sized to enormous creatures in the space of some 5 or 10 million
years. Reaching lengths of 30 to 50 feet, and with bodies that were quite deep
and sturdy, the shastasaurs were the largest ichthyosaurs and among the largest
of the marine animals of the M e s o z o i c . T h e still-unnamed giant ichthyosaur
being excavated by Nicholls might turn out to be a shastasaur. T h e first of the
really large ichthyosaurs was the 33-foot-long Cymbospondylus. It was originally
believed that the tail portions of this species and of the giant shastasaurids
lacked the tailbend that characterized the later ichthyosaurs, but Jennifer
Hogler (1993) reexamined the tail portion of Cymbospondylus and Shonisaurus
and found the wedge-shaped caudal vertebrae that are the hallmark of the
tailbend, suggesting that these early ichthyosaurs had a tail structure similar
to that of the later forms.
Found in the m i d d l e Triassic deposits of Nevada, Cymbospondylus was about
the size of an adult male killer whale, with a short (or perhaps n o ) dorsal fin.
T h e first evidence of the existence of this giant ichthyosaur was a 3-foot-long
skull found by Annie Alexander in Nevada in the early years of the twentieth
century, but afterward, more specimens were found, including an almost
complete skeleton. Cymbospondylus fossils have been found in N o r t h America,
Spitsbergen, and Europe, and recently, two skulls were found in the U p p e r
Triassic Falang Formation in Guizhou province, China, extending its range to
THE
93
The
30-fool-long
ichthyosaur
Cymbospondylus with a
juvenile.
another continent ( L i and You 2002). T h e lower jaw contained teeth only on
Ichthyosaurs
delivered
their
young
the forward part, and it has the smallest orbit (and therefore the smallest eyes)
known
whether
they
Paleontologists do much of their work in the field, but there are occasions
when d i g g i n g through m u s e u m collections also produces some interesting
results. In the collection of the Academy of N a t u r a l Sciences of Philadelphia,
Chris M c G o w a n may have found another ichthyosaur that was as big as Shonisaurus, or maybe even bigger. In 1996, he described a "giant ichthyosaur of the
Early Jurassic," identified from a mislabeled bone in the academy's collection.
Because of its size, it had originally been identified as a coracoid, which is part
of the shoulder girdle and one of the larger parts of any ichthyosaur skeleton.
As he recounts in The Dragon Seekers (2001), M c G o w a n saw that "the robust
element was not part of the shoulder at all but part of the skull, namely a
quadrate bone." A n d what a quadrate bone! " T h e entire animal must have
been colossal larger than any ichthyosaur ever found in England." M c G o w a n
then revisited the collection in the N a t u r a l H i s t o r y M u s e u m in London,
where he found a massive scapula and teeth that were much larger than those
of any known ichthyosaurs. He wrote, "I suspect the massive isolated teeth
belong to the unknown g i a n t . . . as large as Shonisaurus. , . all we know so far
about this enigmatic giant is that it reached lengths upwards of about fifty feet
(15 meters) and that it had massive teeth with sharp edges and short crowns."
M o s t ichthyosaur fossils have been found in Europe and N o r t h America,
with England and Germany as the prime locations. S o m e fragmentary m a t e rial was found in Argentina, but recently, more complete fossils of various
marine reptiles, including pliosaurs, turtles, and ichthyosaurs, have been
found in Neuquen province of the Argentine Andes. As M a r t a Fernandez
(2000) of the M u s e o de La Plata wrote, "a rich fauna of marine reptiles has
been discovered from T i t h o n i a n levels of the Vaca M u e r t a Formation, exposed as several localities in the N e u q u e n Basin." T h e first Argentine ichthyosaurs were referred to Ophthalmosaurus (Gasparini 1985), but one of the
recent specimens has been reassigned to a new genus, Caypullisaurus, which
differs from Ophthalmosaurus in the osteology of the forefin. T h e r e is also a
long-snouted species named Chacaisaurus ( F e r n a n d e z 1994) and another new
species from the Los M o l l e s Formation that was named Mollesaurus (Fernandez 1999). T h e Argentine Ophthalmosaurus from the T i t h o n i a n levels of
Vaca M u e r t a ("dead c o w " ) was about 148 m i l l i o n years old, but Mollesaurus
was from the Bajocian, perhaps 25 m i l l i o n years older.
Investigators continue to uncover more ichthyosaur fossils, often in unexpected locations. For example, when D i n o Frey, M a r i e - C e l i n e Buchy, and
Wolfgang Stinnesbeck examined the material in the collection of the Faculty
of Geosciences in Linares, M e x i c o , they found (but have not yet d e s c r i b e d ) "a
large number of vertebrae and segments of columnae vertebrates of ichthyosaurs . . . but no cranial material." In their 2001 abstract (in which they also
discussed the giant pliosaur known as the " M o n s t e r of A r a m b e r r i " ) , Frey et
al. wrote that they were planning to return to M e x i c o because they "expect to
discover more material in order to determine the taxonomical and paleobiogeographical importance of these assemblages."
In recent years, China has become one of the world's most important
sources for fossils, particularly of the specimens that have led paleontologists
to recognize that many early dinosaurs had feathers, demonstrating a direct
connection between creatures such as Archaeopteryx and today's birds. C h i n a is
1 H t
95
also offering up fossil ichthyosaurs. In 1999, C h u n Li reported on an ichthyosaur fossil from Guizhou province in southwestern China that he named
Qjanicbthyosaums, for Qian, which is another name for Guizhou. T h e n Betsy
Nicholls, Chen Wei, and M a k a t o M a n a b c published a lengthy description of
a complete specimen of Qiankhthyosaurus, also from Guizhou, that had a short
snout; small, closely spaced teeth; and huge eyes. It is believed to be closely
related to Toretocnemus, found in late Triassic deposits in Shasta County,
California, and originally described by M e r r i a m in 1903. " T h e close relationship between Qiankhthyosaurus and Toretocnemus indicates a trans-Pacific d i s tribution of [ichthyosaur] faunas in the late Triassic," wrote Nicholls and her
colleagues in 2003.
In M a y 2000, N a t h a l i e Bardet and M a r t a Fernandez published a description of a new ichthyosaur from the U p p e r Jurassic lithographic shales of
Solnhofen, one of the first new specimens described from the area in almost
50 years. "Indeed," they wrote in their historical review, "with the exception of
Meyer's [1863] skull, kept at London, the ichthyosaur collections from the
lithographic limestones of Bavaria (then housed in the M u n i c h State M u s e u m ) were completely destroyed during W o r l d W a r II." T h e new species is
based on two specimens, one unearthed in 1954 (originally referred to as
Macropterygius posthumus), and another found in 1990 and called Macropterygius
trigonus. T h e type specimen is one that had previously been identified as
Ichthyosaurus leptospondylus ( W a g n e r 1853), but it was destroyed in the bombing
of M u n i c h . Because the two specimens of Macropterygius were found to differ
significantly from known Ichthyosaurus specimens, Bardet and Fernandez proposed that the species be renamed Aegirosaurus leptospondylus, from Aegir, god of
the oceans and seashores in German and Scandinavian mythology. It was a
m e d i u m - s i z e d ichthyosaur, about 6 feet long, with a long, slender snout
packed with small teeth strongly anchored in the jaws. Its eyes were large,
and the sclerotic rings were composed of fourteen overlapping thin plates
(Ophthalmosaurus's eyes had fifteen p l a t e s ) . Impressions of the soft tissue were
preserved all around the body, except on the skull, and showed the typically
lunate tail and four limbs. T h e gracefully curved forelimbs were long and
narrow, and the hind limbs were shorter and broader.
T h e Solnhofen limestone occasionally preserves impressions of the skin,
and in the case of Aegirosaurus, Bardet and Fernandez observed what they
Skeletal
tion
saur O p h t h a l m o -
surprising aspect of all ichthyosaur soft tissue specimens is the lack of preservation of typical reptilian scales," and for the most part, negative evidence
that is, the absence of scales in any known specimens has led most researchers to conclude that the skin of ichthyosaurs was smooth, like that of
dolphins. In an 1853 study, H e n r y Coles described scales for ichthyosaurs
("when grouped together on the surface, they appear like hairs or spines; but
when detached, by their short and generally flattened shapes, they approximate more nearly to scales"), but it is obvious, just by looking at the plate in
Coles's paper, that the "scales" were actually cephalopod hooklets. W h a t was
supposed to indicate what the skin of the ichthyosaur looked like actually
showed what it ate.
In his 1973 discussion of the cranial m o r p h o l o g y of Ichthyosaurus, M c G o w a n
discussed the size of the brain: "By reptilian standards the brain was extensive. . . . T h e cerebellum was very large, a condition which is indicative of a
high level of locomotor integration. . . . T h e corpus striatum is the seat of
innate behavioral activity and it seems reasonable to conclude that the ichthyosaur possessed a wide spectrum of instinctive behavioral patterns." T h e n
he wrote:
From observations of specimens in which the unborn offspring are preserved it is clear that few were born at any one time and these were
delivered from the cloaca tail first. T h e birth would certainly have been
reconstruc-
of the ichthyo-
large
ring,
downturn
the
in the vertebral
column,
profusion
and the
of ribs.
eye
attended by a considerable degree of parental care, and may also have been
accompanied by displays of social co-operation by other individuals. It is
very probable that ichthyosaurs were gregarious and some evidence for this
is available from Germany where aggregations of specimens have been
found in some quarries. T h e gregarious habit is associated with cooperative behavior which is further evidence of cerebral activity at a high
center.*
Based on size, tooth structure, comparison with living animals, and many
other variables, paleontologists can make educated guesses about the diet of a
particular kind of animal. It is rare that fossils yield evidence of the actual
prey items (the fossils of some predatory fishes have been found with the
fossils of undigested smaller species in their stomachs), but some ichthyosaurs
have been found with the fossilized hooks of cephalopods in their gut, giving
a clear indication of what they ate. H o w the ichthyosaurs actually caught their
prey has also been a subject of speculation, and for those with dolphin-like
jaws and teeth, the answer appears obvious: they swam through a school of
fishes or squid and snapped up the smaller animals, much the way many
dolphin species do today. We know that dolphins have an additional weapon
in their arsenal in the form of echolocation; they can emit sounds and listen
to the returning echoes to get a fix on the location, speed, and even type of
prey. C o u l d the ichthyosaurs echolocatc? Probably not. U n l i k e the ears of
dolphins, which are surrounded by a layer of s p o n g y tissue, the middle and
inner ear bones of ichthyosaurs were uninsulated and part of the skull, which
precludes the directional hearing necessary for the analysis of returning
echoes. In his discussion of the cranium of Ichthyosaurus, M c G o w a n (1973a)
also wrote that "the last piece of evidence to support the absence of direction
location is the nature of the sensory receptor. It has already been noted that
the lagena [an extension of the saccule of the ear] was probably a small
structure, and this evidence, slender as it is, is not suggestive of an acute sense
* Several specimens found in the same place does not necessarily show gregarious behavior,
because a number of specimens might have died in the same place but not necessarily at the
same time.
from Dorset, W h i t e a r (1956) believed that she had identified some brown
pigments.
Evidence has recently been uncovered from a clay q u a r r y in Peterborough,
England that might demonstrate that Jurassic ichthyosaurs ate belemnitcs,
and what's more, after they ate them, they vomited up the indigestible guards.
According to paleontologist Peter Doyle of Greenwich University in London, the find consisted of fossilized shells, whereas previous discoveries had
contained only the a r m hooklets. In an interview in New Scientist (Pearce 2002),
Doyle said, "It showed that ichthyosaurs behaved much as sperm whales do
today"; that is, they ate the belemnites and then vomited up the bullet-shaped
shells. H o w did Doyle know they were vomited up? Because the shells showed
distinctive signs of being etched by the ichthyosaurs digestive juices. H o w did
he know they came from ichthyosaurs? Because 160 million years ago, as
shown by other fossils, ichthyosaurs lived there. In an earlier paper, Doyle and
M a c d o n a l d (1993) wrote that " [ b e l e m n i t e ] hooks would effectively interlock
in the stomachs of predators m a k i n g the normal process of excretion difficult
and regurgitation a more likely disposal process." Other paleontologists are
not so sure that these shells were regurgitated, let alone regurgitated by
ichthyosaurs. Also, it has not been shown unequivocally that it was stomach
acids that etched the guards; the m a r k s m i g h t have been made by acidic
sediments in the waters in which the fossils were found.
D o l p h i n s do it, but did ichthyosaurs do it? R i c h a r d Cowen suggested that
ichthyosaurs leaped out of the water, but the question is unanswerable from
the fossil record which is all we have to go on with ichthyosaurs. T h e i r sleek
shapes imitate those of dolphins and argue for leaping ability, as does their
need to remain near the surface in order to breathe.* Also, it has been shown
* Michael Caldwell of the University of Alberta at Edmonton does not believe that
ichthyosaurs should be compared with dolphins. In a November 1 9 9 9 letter to me, he
wrote, "There is nothing dolphin-like about an ichthyosaur. However, there is something
ichthyosaur-Iikc about a dolphin. . . . Ichthyosaurs did metabolic things that some extant
reptiles do some more or less closely related (crocodiles and marine iguanas) and unrelated (sea turtles) all of which, like sharks and tunas, accomplish their active marine habits
in ways that are mctabolically unfamiliar to metabolic solutions of mammals. There are
similarities, but each similarity represents cither ancient common ancestry (they are all
vertebrates) or convergence (fins in tuna, ichthyosaurs, and dolphins). I see no imperative to
explain some unique mechanism that was mammalian-like. Rather, to simply indicate that
mammals have neither a superior nor inferior metabolism and physiology. That many
animals work with non-mammal systems to accomplish what mammals accomplish. The
marvel is not in the mammal-hke condition, but rather in the diverse physiologies that
achieve similar results. Kind of a 'there's more than one way to skin a cat' sort of thing."
We will probably never know if ichthyosaurs leaped out of the water for
greater s w i m m i n g efficiency, for more effective fish capture, or just for fun,
but the image of a fish lizard flying through the air is an intriguing one.
S w i m m i n g with fins and tail seems a simple enough business: flexing the
tail against water resistance provides forward movement, and the fins act as
planes to adjust the angle. But it is not nearly that simple, and opinions differ
on the properties of the ichthyosaur tail, with the vertebral column in the
lower rather than the upper lobe as it is in sharks. W h e n Keith T h o m p s o n
and David S i m a n c k (1977) studied locomotion in sharks, they concluded that
the heterocercal (one lobe longer than the o t h e r ) caudal fin is also instrumental in changing the pitch of the shark's body in the water, and subtle adjustments enable the shark to turn and accelerate quickly. Because the ichthyosaurs had to come to the surface regularly to breathe, they had another set
of problems to solve in addition to s w i m m i n g through the water: they had to
compensate for the downward thrust generated by the tail fin. Michael Taylor
(1987b) wrote that "the caudal fin of ichthyosaurs is usually assumed to have
the p r i m a r y function of propelling the animal, but this does not explain why
many ichthyosaurs had a caudal fin of the reversed heterocercal type." He
suggested that the tail fin alone could compensate for residual buoyancy, and
"there would be no need for the pectoral and pelvic fins to produce any lift."
In 1992, M c G o w a n wrote a paper in which he contended that simply reversing
the tail structure did not mean that the tails of sharks and ichthyosaurs
functioned in the same way, just because the bones were in one lobe rather
than the other. " T h e two structures," he wrote, "are not strictly analogous,
and there are functional g r o u n d s why the ichthyosaurian tail should not
generate vertical forces." He then suggested that even though they operate in a
horizontal plane, the symmetrical tail flukes of cetaceans might make a better
analogue for the tail fin of ichthyosaurs. Because no human has ever seen an
ichthyosaur, our discussions about the dynamics of its s w i m m i n g only point
up the problems inherent in interpreting fossil animals.*
0
ichthyosaurs did not use their tails for locomotion but rather "flew" underwater like
penguins, using their foreflippers for propulsion. In his third-year dissertation at the
University of Southampton ( U K ) , Darren Naish refuted this peculiar notion, and in a
popular article in 1 9 9 8 , he wrote, "One clear correlation that docs appear to be true tor
swimming vertebrates concerns the tail. Essentially, if an animal has a propulsive surface on
could probably not have escaped being inertial h o m e o t h e r m s and they probably maintained b o d y temperatures greater than that of the surrounding sea."
Size or similarity of body plans does not automatically confer endothermy on
ichthyosaurs, but such comparisons are steps in the direction of understanding how these marine reptiles functioned. Unfortunately, other extinct large
marine reptiles d i d not resemble tunas or sharks; some looked like crocodiles,
others like s w i m m i n g sauropod dinosaurs, and some like no other creatures
before or since. H o w did marine crocodiles, plesiosaurs, and mosasaurs manage to retain enough body heat to chase and capture their prey in the ocean?
In a 1993 article about predatory marine reptiles, Bakker described the ichthyosaur Baptanodon (a synonym for Ophthalmosaurus) as "having the deadly
advantage of slashing speed. T h e i r bodies have the 40-knot shape preferred
by evolution for all its fastest s w i m m i n g creations." Nowhere in this article
does he address the thorny question of how an ichthyosaur might have
achieved this "slashing speed." T h i s is a curious omission for Bakker, who is a
champion of the idea that ancient predators (theropod d i n o s a u r s ) had to
have been w a r m - b l o o d e d .
In a 1996 discussion of Chensaurus ( w i t h Crippta and Utatsusaurus, the earliest
known ichthyosaur), M o t a n i , You, and M c G o w a n described this genus as
long and slender (the fossil is about 30 inches l o n g ) , with a very high vertebral
count that suggests an anguilliform ( e e l - l i k e ) s w i m m i n g motion, probably
similar to that of some scyliorhinid sharks. Later ichthyosaurs developed a
body plan that was carangiform (shaped like a jacklike fish) and finally
thunniform (shaped like a t u n a ) . By the time of the Triassic Shonisaurus,
s w i m m i n g efficiency had been sacrificed for sheer size.
Early paleontologists recognized two forms of ichthyosaurs: a broadfinned type they called latipinnate, from the Latin latus, meaning "broad," and
pinna, for "fin"; and longipinnate, from longus, which means "long." As earlier
defined, latipinnate ichthyosaurs tended to be moderate in size, not exceeding
12 to 13 feet in length, but some of the longipinnatcs were gigantic; Cymbospondylus reached a length of 33 feet, and Shonisaurus was 50 feet long, bigger than an
adult gray whale. A l t h o u g h the two types of ichthyosaurs occupied the same
waters, the latipinnates became extinct by the late Jurassic, while the longipinnates lasted another 40 million years, into the late Cretaceous. Although it
of
Ichthyosaurus.
Instead of the
familiar five
the
developed
hones
fingers,
ichthyosaurs
tile-shaped
arranged on
digits,
of tile-shaped bones arranged on the long axis as digits, but squashed to-
hut squashed
formed
saurs with three digits per flipper; a couple with four; many with five; and
flipper-shaped
appendage.
the Lower Liassic of England, have 27 elements in the longest digit, and W a d e
(1984) counted 30 in the flipper of Platypterygius.
N o w considered by some to be the same genus as the British Temnodontosaurus ( M c G o w a n 1995c), Leptopterygius ("narrow fin") was a fast-swimming
predator of the Jurassic seas. (Leptopterygius has been renamed Leptonectesf) Temnodontosaurus reached a length of 29 feet and d i d not share the toothlessness of
Stenopterygius. It was as large as a modern killer whale, but its teeth were smaller.
Its eyes were as large as or larger than those of Ophthalmosaurus, and like many
solid,
The
iO-fiot-lont
early
Jurassic
ichthyosaur
the largest
of any
animal
seems
was only about two-thirds as large as the gigantic Shonisaurus and had teeth in
reasonable
to
proportion to its size, suggesting that it fed on large prey items, probably
assume thai it
hunted in
reduced
light conditions.
dramatic
black
while pattern
purely
The
and
is
conjectural.
The
100-miUion-
year-old
ichthyosaur
Eurhinosaurus
(top) bears a
remarkable
resemblance to the
modern
swordfish
Xiphias gladius.
(Inlike
fish,
the sword-
the
had teeth.
ichthyosaur
Earl Herald (who ought to have known better) wrote, "the sword may be used to impale
fishes during feeding," which seems highly unlikely, because the prey fish would offer no
resistance to the impaler, and even if such a process could work, the swordfish would be
unable to get at the dead fishes stuck on the end of its nose.
way through schools of fishes and gathering up the incapacitated victims, but
inherent problems make direct observations virtually impossible.") M o s t ichthyosaurs had teeth, and they u n d o u b t e d l y used them to catch their prey. It is
not clear, however, what purpose was served by the teeth in that part of the
upper jaw of Eurhinosaurus. Because of the severe overbite, the upper teeth
could not make contact with the teeth in the lower jaw or, for that matter,
with anything at all.
W h i l e excavating a drainage channel near Stowbridge in Norfolk, England,
in 1958, workmen came upon a large, long-snouted ichthyosaur skull, along
with some vertebrae, ribs, and other fragments. It was provisionally identified
as a species of the Jurassic ichthyosaur Ophthalmosaurus, but the eye sockets
were not nearly as large, so it was reclassified in 1976 by M c G o w a n . Because it
had powerful jaws and teeth, he n a m e d this killer whalesized ichthyosaur
Crendelius mordax the generic name from Grendel, a legendary monster in
Beowulf, and the specific name from mordax, Latin for "biting." A l o n g with
Excalihosaurus, Crendelius was one of McGowan's more inspired names, but alas,
it didn't endure. In 1989, another specimen was found in the Kimeridge C l a y
of Dorset, and when M c G o w a n examined the new material ( p a r t i c u l a r l y the
newly excavated forefin), he realized that it should actually be placed in an
existing genus (Brachypterygius) that had been erected in 1922 by German paleontologist Frederich von H u e n e . Too bad; Grendelius was a wonderful name.
M a n y of the ichthyosaurs had large eyes, but the largest relative to b o d y
size belonged to the appropriately n a m e d Ophthalmosaurus; its eyes were 8
inches across in a 15-foot-long animal. ( T h e eyes of a 15-foot pilot whale are
about an inch across, and the eyes of a 100-foot blue whale, probably the
largest animal that has ever lived, are about 8 inches across.) In a 1999 study,
M o t a n i , Rothschild, and W a h l measured the sclerotic rings of Temnodontosaurus and found that a 30-foot-long individual had an eye that measured
nearly 10 inches in diameter, m a k i n g it the "largest eye in the history of life."
But Ophthalmosaurus had the largest eyes relative to b o d y length of any vertebrate that has ever lived.* T h e authors found that the f-number (the measure
* T h e giant squid
(Arcbiteutbis)
often described as being "as big as dinner plates" (a standard dinner plate is 10 inches in
THE
] CMTH J 0 5 A
7
UliS
109
of relative aperture used in camera lenses) for Ophthalmosaurus was the lowest
the
of any of the ichthyosaurs (the lower the f-number, the more light the eye
late Jurassic
ichthyosaur
the much
larger Tcmno-
picks u p ) . T h e y believe that this ichthyosaur had the ability to sec where light
barely penetrated and that it "probably had higher visual sensitivity than a
cat," an animal justly renowned for its vision in low light. We know that these
big-eyed reptiles were habitual deep divers, because their fossilized skeletons
showed evidence of the bends. We have no way of knowing if the squid that
dontosaurus had
species are, it seems a reasonable assumption, and big eyes with a very low
the size of
Ophthalmosaurus.
" W h y did some ichthyosaurs have such large eyes?" asked Stuart H u m phries and Graeme Ruxton in a 2002 article in the Journal of Experimental Biology.
T h e y answered, "sensitivity to low light at great depth has recently been
diameter) or even "automobile hubcaps," but the scientific literature contains no such
dimensions (Ellis 1 9 9 9 ) . M o t a n i et al. ( 1 9 9 9 ) cited Roper and Boss's 1 9 8 2 Scientific American
article, which states that the eyes of the giant squid are "enormous, larger than the headlights of an automobile . . . with a diameter approaching 25 centimeters ( 1 0 inches) they arc
the largest eyes in the animal kingdom."
] CH'TM i O 5 A t
1
US
in
where they are painful but not life-threatening. T h a t ichthyosaurs got the
bends suggests that they had not perfected their diving physiology the way
today's marine m a m m a l s have. ( R o t h s c h i l d and M a r t i n have also found evidence of the bends in deep-diving mosasaurs.)
T h e eyes of Ophthalmosaurus may have been among the largest relative to
body size (those of Tenmodontosaurus were larger in absolute size), but how
these big-eyed ichthyosaurs captured their food is not immediately evident.
One m i g h t assume that a creature built like a dolphin would have a similar
diet, but most of the known fossil skulls of Ophthalmosaurus are toothless, a
condition called cdeutulousness. T h i s has led to an assumption ( A n d r e w s 1910)
that Ophthalmosaurus must have fed on soft-bodied animals like squid and may
have used some sort of suction m e t h o d to capture them. But when Angela
Kirton (1983) examined the skulls of various Jurassic ichthyosaurs from England, she found that many of the large skulls did indeed have teeth. T h e r e are
two ways of explaining edentulousness in adult Ophthalmosaurus: either they
The
iz-Joot-long
ichthyosaur
Platypterygius
shown chasing an
ammonite,
extinct
an
cephalopod.
lost their teeth as they matured, or they had perfectly serviceable teeth that
were loosely attached (like the teeth of some s h a r k s ) and were lost during the
hundreds of millions of years between the time the animal died and the time
the fossils were discovered. S o m e believe the former explanation, and others
subscribe to the second theory. Ryosukc M o t a n i , who provided this infor-
JT
U'R_S
113
T h e "youngest" ichthyosaur fossils (so far) are about 93.5 million years old
and were found in the U p p e r C e n o m a n i a n of Bavaria (Bardet et al. 1994).
Platypterygius was a large animal, reaching a length of 23 feet. Its jaws were long
and slender, its eyes were relatively small (but only when compared with its
huge-eyed relatives such as Ophthalmosaurus), and its forcflippcrs were the
widest of any ichthyosaurs (Platypterygius means "broad fin"). T h i s species was
also polydactylic, with as many as ten rows of squarish bones in the flipper.
Sander (2000) wrote, "this is the only instance of polydactyly in amniotes that
is not pathologic." Like all later ichthyosaurs, this genus had a symmetrical
tail fin, with the vertebral support in the lower lobe. T h e tail was certainly
used for propulsion, but the narrow-based fins were probably used for sculling, adding another element to the propulsive powers of this reptile. ( T h e
reduced hind fins were probably not particularly useful and are comparable to
the paired pelvic fins of sharks, which probably function as secondary stabilizers in s w i m m i n g . )
Several attempts have been made to place ichthyosaurs in deposits later
than the C e n o m a n i a n (93.5 m i l l i o n years a g o ) , but these have proved unsuccessful; in her 1992 review of the stratigraphic evidence, Nathalie Bardet
wrote, "A review of these post-Cenomanian ichthyosaur remains reveals that
all are doubtful either from a stratigraphic or systematic point of view."
After listing the various "systematically doubtful specimens" and the "stratigraphically doubtful specimens," Bardet speculated on the extinction of the
ichthyosaurs:
T h e group, being pelagic since the Jurassic, was potentially insensitive to
the great anoxic event characterizing this period. Biological factors such as
replacement by mosasaurs arc debatable, as these two groups probably did
not occupy the same ecological niche. T h e great extinction suffered by
marine invertebrates, especially cephalopods, at the Ccnomanian-Turonian
boundary, may have preferentially affected specialized predators such as
ichthyosaurs more than generalist ones such as plesiosaurs, in terms of a
break in their food chain. If the extinction of ichthyosaurs is linked to the
Cenomanian-Turonian extinction events, such a break in the food chain
may be proposed as an extinction scenario.
evolutionary paradigm, viz., that by the end of the J u r a s s i c / e a r l y Cretaceous, ichthyosaur m o n o p o l y of a thunniform body shape and commensurate mechanical design ended and that this heralded their demise. T h e
emergence of fast-swimming fishes with streamlined bodies placed new
energetic costs on ichthyosaur prcdation from the perspectives of both
more evasive prey and more effective predators. Furthermore, ichthyosaurs
were K-stratcgists, producing a small number of large young. Predator
avoidance in ichthyosaur juveniles, by high-speed Sight, was placed under
greater stress from the newly emerging, streamlined, fast-swimming bony
fishes and sharks. . . . By the closing stages of the Late Cretaceous pursuit
prcdation in marine reptiles was almost completely replaced by lie-in-wait
or ambush predatory tactics. T h e hydrodynamic body shape of ichthyosaurs gave way to the long-bodied design, exemplified in the mosasaurs, a
group of marine reptiles that arose in the latest stages of the Cretaceous.
T h e latter design is more effective for rapid starts and burst speeds, useful
in ambush prcdation. A m o n g plesiosaurs, only the less hydrodynamic
long-necked elasmosaurs survived to the last stages of the U p p e r Cretaceous. T h e i r extremely long necks, rather than long bodies, may have been
an alternative adaptation for ambush strikes.
In a 1999 article devoted to the aberrant African mosasaur Corcnyosaurus,
Lingham-Soliar speculated on the extinction of the ichthyosaurs:
An understanding of mosasaur evolutionary success is critical to our understanding of the extinction and reduction of other marine vertebrates
during the Cretaceous. Jurassic and Cretaceous ichthyosaurs had torpedoshaped bodies and a deep, hydrodynamically-shapcd tail, resulting in highspeed swimming. It is almost paradoxical that they should have given way
to the hydrodynamically inferior mosasaurs. It points, however, to a chang-
ing environment that was clearly conducive to long-bodied animals. Jurassic ichthyosaurs had become "boxed in" by too much specialization. H i g h speed, highly evasive fishes were becoming dominant in Cretaceous waters
and pursuit predatory tactics of ichthyosaurs, for instance, are energetically
very expensive. . . . T h e sit-and-wait or ambush strategies favored . . .
mosasaurs and it is no coincidence that Cretaceous ichthyosaurs were
reverting to longer forms such as first seen in their early history in the
Triassic. But it was too little and too late for them. It is even less of a
coincidence that at precisely this time the ichthyosaurs became extinct and
the supreme long-bodied mosasaurs arose.
T h e ichthyosaurs are a m o n g the most highly developed reptiles that ever
lived. T h e y were superbly designed for earning a living, with their toothstudded beaks, large eyes, powerful tails, and two sets of steering fins. T h e y
came in all shapes and sizes, from small to extra large. S o m e of them might
have been endothermic. T h e y bypassed the limitations imposed by having to
lay eggs on land by giving birth to live young. T h e y were to the oceans what
the ( n o n a v i a n ) carnivorous dinosaurs were to the land, and for the same
amount of time. W h a t happened to them? We can only speculate. "Well
before the close of the Jurassic," wrote M c G o w a n (1973a), "their numbers
began to dwindle, and relatively few [species] survived into the Cretaceous.
W h i l e the last chapters of the Age of Reptiles were being written on the land,
the last of the ichthyosaurs slipped quietly and unpretentiously into oblivion."
The Plesiosaurs
There were no real Sea Serpents in the Mesozoic Era, but the Plesiosaurs were the next thing
to it. The Plesiosaurs were reptiles who had gone back to the water because it seemed like a good
idea at the time. As they knew little or nothing about swimming, they rowed themselves around
in the water with their Jour paddles, instead of using their tails Jor propulsion like the brighter
marine animals. This made them too slow to catch fish, so they kept adding vertebrae to their
necks until their necks were longer than all the rest of their body. Then they would dart their
heads at the fish from a distance of twenty-five or thirty jeet. Thus the Plesiosaurs resembled the
modern Sea Serpent above the water-line, though they were almost a total loss farther down.
They might have had a useful career as Sea Serpents, but they were before their time. There was
nobody to scare except fish, and that was hardly worth while. Their heart was not in the work.
As they were made so poorly, Plesiosaurs had very little fun. They had to go ashore to lay their
eggs and that sort of thing.
swallowing pebbles after each meal to grind their food. At least, pebbles have been found near
fossil Plesiosaurs, and to a scientist that means that the Plesiosaur had a gizzard. During
the Cretaceous Period many of the inland seas dried up, leaving the
Plesiosaurs stranded without any fish. Just about that time
Mother Nature scrapped the whole Age of Reptiles
and called for a new deal.
see what she got.
Will Cuppy (1941)
And you
Less enjoyable but probably more useful than Cuppy's definition is Glenn
Storrs's 1997 classification of the genus Plesiosaurus, one of the many genera
that make up this fascinating group:
Plesiosaurs ( D i a p s i d a : Sauropterygia: Plcsiosauria) arc extinct M e s o z o i c
marine reptiles comprising one of the most successful and widely d i s tributed groups of marine reptiles. T h e y developed a wide range early in
their history and some representatives of the cladc survived into the
Maastrichtian, becoming extinct perhaps only at the terminal Cretaceous.
T h e evolutionary and systematic relationships of the Plcsiosauria, however, are almost completely unknown. T h e group appears as isolated bones
and associated partial skeletons in the M i d d l e Triassic ( A n i s i a n ) of Germany, but the first unambiguous, fully articulated specimens occur in the
uppermost Triassic and Lower Jurassic ( L i a s s i c ) of England. By Liassic
times, the plesiosaurs were particularly diverse, already fully marine, and
highly modified from the presumed terrestrial condition of their forebears.
T h e name plcsiosaur means "near reptile," and it was bestowed on the earliest
fossils by Conybeare because he believed that they represented animals that
were on their way out of the sea to become the terrestrial reptiles.* We now
know that it was actually the reverse they are descended from land reptiles
that returned to the sea. T h e s e great oceangoing creatures dominated M e s o zoic seas and were powerful predators, armed with a mouthful of sharp teeth.
Unlike the ichthyosaurs, which looked rather like dolphins, plesiosaurs came
in all shapes and sizes, many of them unique. S o m e had short necks and huge
heads, while others had long necks and tiny heads. All four limbs were
modified into flippers, and the shoulders and pelvic girdle were formed of
broad sheets of bone to which the powerful s w i m m i n g muscles were attached.
* On the subject of the name, Robin O'Keefe ( 2 0 0 2 a ) wrote: "The term 'plcsiosaur,'
meaning 'near lizard' is not an informative name from a modern perspective. However,
when Conybeare ( 1 8 2 2 ) coined the term to describe fossils from the English Lias, little was
known concerning any extinct marine reptile. The realization that plesiosaurs were a
completely extinct group was significant at a time when the occurrence of extinction itself
was uncertain. 1 hesc 'near-reptiles' were named at a time when there was no need, and no
context, for a more specific term.''
T h e flippers were long and narrow, but there is no consensus among paleontologists as to how these great reptiles actually moved through the water. Did
they pull through the water in a "breaststroke," or "fly" like turtles or penguins? T h e r e are no fossil plesiosaurs that contain unborn fetuses, so we do
not know if these reptiles gave birth to live young in the water or came ashore
to lay their eggs. T h e dense rib cage, especially in the belly region, has
suggested to some that they might have come ashore, but with flippers instead
of feet and, in some cases, an extremely long neck, they would have been very
awkward out of the water and susceptible to prcdation by land animals such
as crocodiles or carnivorous dinosaurs. It now seems unlikely that the plesiosaurs ever left the water.
T h e description of a plesiosaur as "a snake threaded through the body of a
turtle" has variously been attributed to Convbeare, De la Bechc, M a n t e l l ,
Owen, and both W i l l i a m Buckland and his son Frank, but its actual origin
remains a mystery. In his 1824 description of the first plesiosaur fossil, C o n y beare wrote, "In its motion, the animal resembled the turtle more than any
other, and the turtle also, as it was better remarked, could we divest it of its
shelly case, would present some slight approach in its general appearance to
the plesiosaurus." In 1837, M a n t e l l wrote, " T h e reptile combines in its structure the head of a lizard with teeth like those of a crocodile, a neck resembling
the body of serpent, a trunk and tail resembling of the proportions of a
quadruped, with paddles like those of turtles." In his 1914 Water Reptiles of the Past
and Present, Samuel W i l l i s t o n wrote, "It was Dean Buckland who facetiously
likened the plesiosaurs to a snake threaded through the shell of a turtle, but
what Buckland actually wrote (in the 1836 Bridgcwater Treatise), was: 'To the head
of a lizard, it united the teeth of a crocodile; a neck of e n o r m o u s length,
resembling the body of a serpent; a trunk and tail having the proportions of
any ordinary quadruped, the ribs of a chameleon and the paddles of a whale.' "
T h e enigmatic Pistosaurus, which gets its name from the Greek pistos, for
"liquid," the medium in which it lived, was found in the M u s c h c l k a l k ( m i d d l e
Triassic) of Germany. It is known only from a skull and a postcranial skeleton
that was found in the same location, but not in direct articulation ( S u e s
1987b). "If this association is correct," wrote Carroll (1988), "this genus
combines a postcranial skeleton like that of the typical nothosaurs with a
longer-
elasmosaurs,
Muracnosaurus
gels its name from
muraena, the l a t i n
namefor
the
moray
reptile.
Muracnosaurus
was 10 feet long.
This is
Plesiosaurus
dolichodeirus,
described in 1811 by
Henry Dc la
and
Conybeare
long powerful jaws could have bit his neck in two without ceremony."
skeleton of a
from
Mary
Lyme Regis.
creature Plesiosaurus dolichodcirus, which means "near lizard with a long neck."
T h e skeleton, now in the Natural H i s t o r y M u s e u m in London, was 7V2 feet
long from the tip of its skull to the tip of its tail. Its little head and long neck
were approximately the same length as its torso and tail, and it had needle-like
teeth in both the upper and lower jaws. S o m e recent books indicate incorrectly that Plesiosaurus means "ribbon lizard," supposedly for the ribbon-like
the
"fossil
animal" found
Beche
William
by
Aiming at
The
History
in
London,
is ~)Vijeet long.
appearance of the neck spines along its backbone, but there is no basis in
Greek or Latin for such a reading. Conybeare wrote, " T h e name I have
originally given to this animal, Plesiosaurus (approximate to the Saurians), may
appear rather vague in this stage of our knowledge, and an appellation derived
from its peculiar length of neck might be preferred; but for the present I shall
retain the old generic name, adding for specific distinction the well-known
H o m e r i c epithet Dolichodeirus ("long-necked"), as characterizing the most
striking peculiarity of its osteology." T h e archetypal genus Plesiosaurus (De la
Beche and Conybeare 1821) became a sort of g r a b bag for subsequently
discovered plesiosaurs, even though many of them were dramatically different
( S t o r r s 1997).
Joseph Pentland, who was so helpful to Conybeare and Buckland in the
proper identification of the ichthyosaur, also contributed to their understanding of the plesiosaurs. In a letter to Buckland dated June 20, 1820
(reproduced in Delair and Sarjeant 1975), Pentland wrote, "As to M r . Conybeare's new A n i m a l s , I will not pretend to judge, but the disposition of the
bones of the a r m seem to put beyond a doubt that it is very different from the
Ictyosaurus [sic]. . . . From the sketch, I clearly see that the bone can only be
the C o r a c o i d . . . so if you have any influence with M r . Conybeare you would
do well to suggest to h i m to correct the fault he is about to commit by calling
it the Clavicle. . . . T h e name of Plesiosaurus is a very good name I think,
perhaps a little too relative; would it not be better to give some other name
which would express either some peculiar structure in the animal, or one
relative to its high antiquity, while retaining the termination Saurus, which I
think has been very happily chosen." Conybeare seems to have been impressed
with Pentland's suggestions about nomenclature; although he stuck to the
name Plesiosaurus, he added dolichodeirus ("long n e c k " ) to the more complete
specimen found by M a r y A n n i n g in 1824.
In 1832, Dr. R i c h a r d H a r l a n was presented with several enormous vertebrae
by a gentleman named Bry, who had found them on the Ouchita River in
Louisiana. H a r l a n submitted a description of the bones to the American
Philosophical Society in Philadelphia, and his description ("Notice of the
Fossil Bones Found in the Tertiary Formation of the State of L o u i s i a n a " ) was
identified by
1834 (Mosasaurus,
Geosaurus,
Megalosaurus,
Iguanodon,
Ichthyo-
saurus, and Plesiosaurus) and concluded that his bones belonged to a plesiosaur,
80 to 100 feet in length. Accordingly, he n a m e d it Basilosaurus, "king of the
lizards."
It wasn't actually a plesiosaur in fact, it wasn't a reptile at all and it
wasn't 100 feet long. Eight years later, H a r l a n brought the bones to England
and showed them to Richard Owen, who compared the double teeth of Basilosaurus with those of other reptiles and m a m m a l s and concluded that the teeth
were actually those of a primitive whale. Because it wasn't a lizard, the name
Basilosaurus was completely inappropriate, so O w e n proposed the name Zeuglodon, which means "yoked teeth." In the 1842 Geological S o c i e t y report on
Owen's observations, we read: " M r . Owen, in compliance with the suggestion
of Dr. Harlan, who, having compared with M r . Owen the microscopic structure of the teeth of the Basilosaurus with those of the D u g o n g and other
animals, admits the correctness of the inferences of its mammiferous nature,
proposes to substitute for the name of the Basilosaurus that of Zeuglodon,
suggested by the form of the posterior molars which resembled two teeth tied
or yoked together." Unfortunately, the rules of zoological nomenclature d i c tate that the first name has priority, no matter how m i s g u i d e d , so because
H a r l a n named it first, the early whale is still known technically as "king of the
lizards."
Genuine plesiosaur fossils, a m o n g the first of the marine reptiles to be
discovered in England, were highly prized by m u s e u m s and collectors and
often c o m m a n d e d high prices. W h e n a fossilized skeleton of Plesiosaurus
EXHIBITING
DAILY,
SPLENDID
AND
VERY
"PLESIOSAURUS
VALUABLE
FOSSIL
DOLOCHODEIRUS"
T h e "splendid and very valuable fossil" soon became the subject of a bitter
dispute between the m u s e u m s at W h i t b y and C a m b r i d g e , and after many
acrimonious letters between the two institutions, it was finally bought by the
Fitzwilliam M u s e u m of C a m b r i d g e for 230 ( O s b o r n e 1998). It is an almost
complete skeleton, 15 feet from the tip of the snout to the end of the tail, and
its "paddles" were so large that Owen originally named it Plesiosaurus grandipinnins. It can be seen today in the S e d g w i c k M u s e u m of Geology, C a m b r i d g e .
In America, nothing did more to publicize the b u d d i n g science of vertebrate paleontology than the feud between Edward Drinker C o p e and
Othniel Charles M a r s h . In 1868, they were collecting fossils in western Kansas, digging on their own, but also e m p l o y i n g various "collectors" to find
material and ship it back east to them. ( C o p e was affiliated with the A c a d e m y
of Natural Sciences of Philadelphia, M a r s h with Yale University.) In 1867,
T h e o p h i l u s Turner, a physician at Fort Wallace, Kansas, collected three
vertebrae and sent them to Cope, who asked Turner to collect the rest of the
fossil and ship it to Philadelphia. W h e n C o p e examined the bones, he
realized that the original owner was obviously related to the plesiosaurs, but
in addition to its short neck and unusually long tail, there was something
strange about the vertebrae. In the 1869 Proceedings of the Boston Society of Natural
History, Cope published descriptions of various reptiles, including one that he
named Elastnosaurus platyurus ("flat-tailed, thin-plate reptile"). Because most
plesiosaurs had long necks and short tails, he erected a new order to accommodate it, which he called Streptosauria, from the Greek streptos, which means
"turned" or "reversed," referring to the "articular processes of the vertebrae
[which are] reversed in their direction, viz., the anterior looking downwards,
the posterior upwards." Unfortunately, C o p e had completely misunderstood
the vertebrae, assembled the skeleton backward, and put the head at the
wrong end. He was embarrassed into correcting his blunder when M a r s h
gleefully pointed it out, exacerbating the bitter rivalry between the two paleontologists that would last until C o p e died in 1897.
Did
the
plesiosaurs,
long-necked
such
as
In a letter to the New York Herald in 1890, M a r s h described the moment that
he pointed out Cope's mistake:
Elasmosaurus
platyurus, lie on
the bottom,
weighted
stones
they
had
swallowed,
and
reach
upward?
down
with
to
foodjrom
surface?
snatch
the
that he had the whole thing wrong end foremost. H i s indignation was
great, and he asserted in strong language that he had studied the animal for
many months and ought at least to know one end from the other. It seems
he did not, for Professor Lcidy in his quiet way took the last vertebra from
the end of the tail, as C o p e had placed it, and found it to be the atlas and
axis, with the occipital condyle of the skull in position.
C o p e corrected the mistake when M a r s h identified it, but he first claimed that
he had never made it and then said that Leidy was responsible. He tried to
buy up all the copies of the original publication, but a few have survived that
show the head and tail reversed and the limbs on backwards.*
W i t h the head on the right end, Elasmosaurus platyurus now hangs from the
ceiling of the Inland Sea Exhibit at the Academy of N a t u r a l Sciences of Phila* At a meeting of the Academy of Natural Sciences on March 8, 1 8 7 0 , Leidy discussed
delphia, and visitors marvel at its impossibly long neck, short tail, and massive
flippers, which look much too big, even for an animal that was 40 feet long. It
looks for all the world as if another m u s e u m managed to find a fossil of the
Loch N e s s monster. A pair of c o m p a r a b l e skeletons s w i m high above the entrance hall of the Denver M u s e u m of N a t u r a l History, but these are of the
elasmosaur Thalassomedon hanningtoni, from the late Cretaceous of Baca County,
Colorado. In 1970, S. P. W e l l e s published a brief note entitled " T h e Longest
Neck in the Ocean" in the M u s e u m N o t e s of the University of N e b r a s k a
State M u s e u m , in which he discussed the discovery of an elasmosaur in the
Graneros Shale of eastern N e b r a s k a in 1964. It was another 40-foot-long Thalassomedon hannmgloni, nearly as long as Cope's Elasmosaurus platyurus.
In their haste to outcollcct and outpublish each other, C o p e and M a r s h
made a m u d d l e of the identification and classification of plesiosaurs.* As
Glenn Storrs (1999) wrote, "At the most general level, however, the t a x o n o m i c
status of many plesiosaurian taxa remains chaotic, and this particularly a p plies to the several species that have been described from the N i o b r a r a C h a l k .
Niobrara specimens are u n c o m m o n and are often incomplete and the historical holotypcs of such early workers as C o p e and M a r s h arc notoriously so."
Cope's mistakes ("Professor Cope has described the skeleton in a reversed position to the
true one"), but at the same time, he reflected on the modus vivendi of Elasmosaurus platyurus:
"We may imagine this extraordinary creature, with its turtle-like body, paddling about, at
one moment darting its head a distance of upwards of twenty feet into the depths of the sea
after its fish prey, at another into the air after some feathered or other winged reptile, or
perhaps when near shore, even reaching so far as to seize by the throat some biped dinosaur."
* Cope and Marsh were not only rivals for collecting and naming fossils but also albeit
inadvertently were competitors for the title of greatest blunderer in paleontological history. Cope had put the head of Elasmosaurus on the wrong end, but Marsh took the prize, for
he put the wrong head on a dinosaur, a mistake that was not corrected for a century. W h e n
Marsh collected the bones of the huge sauropod Apatosaurus in the Como Bluffs region of
Wyoming in 1 8 8 0 , the head was lacking, so he simply used the head of another gigantic
sauropod named Camarasaurus, which he had found 4 miles away from the body. Although
some people recognized the mistake as early as 1 9 1 5 , it was not corrected until 1 9 7 9 , when
the Carnegie Museum in Pittsburgh replaced the head on its Apatosaurus; shortly thereafter,
the American Museum of Natural History followed suit.
Setting out to rectify this situation, Storrs (1999) and Carpenter (1999)
independently examined the holotypes of all the plesiosaurs described from
the N i o b r a r a C h a l k and attempted to clean up the mess. Of the nine named
species, they found that only three were valid: Polycotylus latipinnis (named by
C o p e in 1869), Styxosaurus snowii (named by W i l l i s t o n in 1890), and Dolichorhynchops osborni (named by W i l l i s t o n in 1902). T h e remainder of the Niobrara
plesiosaurs were synonymized with species from other localities or relegated
to the category of nomina dubia (doubtful n a m e s ) . T h i s is all very complicated, but it points up the state of plcsiosaur taxonomy in the nineteenth
century. T h e s e gigantic sea creatures (identifiable by paddles instead of feet)
had only recently been discovered, and the understanding of their taxonomic
differences was vague. It was little wonder that C o p e and M a r s h identified too
m a n y animals or confused one specimen with another. T h e y were working
with a fossil fauna the likes of which had never before been seen on Earth.
Since Cope and M a r s h so m u d d l e d their identifications, other plesiosaurs
have been found in the American West. Welles and Bump (1949) excavated
the fossil remains of a long-necked elasmosaur that they named Alzadasaurus
pembtrtoni, for R a l p h Pcmberton, M.D., who found the fossil in South Dakota.
( T h e name Alzadasaurus comes from the town of Alzada in southeastern
M o n t a n a , where the holotypc had been found.) In Carpenter's 1999 revision
of the N o r t h American elasmosaurs, he concluded that there were "only five
valid genera
and
Hydralmosaurus serpenlinus,
Thalassomedon
banningtoni.
Li-
Alzadasaurus kan-
// is almost
impossible to picture
Thalassomedon
moving through the
water. The
incredibly long neck
must have acted like
a rudder on the
Jront
of the animal
as it paddled through
the water,
requiring
constant course
corrections.
Woolungasaurus glendoweren-
sis, n a m e d for W o o l u n g a , a reptile in Aboriginal mythology, and the Glendower Station in Queensland, where the fossil was found in 1982. At an
estimated length of 30 feet, it was smaller than Elasmosaurus but similarly
proportioned, with a neck as long as the body and tail combined. One
specimen was found with its skull showing signs of having been bitten by
Kronosaurus, the giant pliosaur that was the scourge of Australian Cretaceous
seas ( T h u l b o r n and Turner 1993). In 1997, Woolungasaurus appeared on an
Australian postage stamp, splendidly tricked out in a suit of black and yellow,
a product of the artist's imagination. No matter what else we have learned
about the extinct marine reptiles, their coloration remains largely a mystery.
A l t h o u g h the plesiosaurs have been known to science and the public for
almost two centuries ( D e la Beche and Conybeare published the first Plesiosaurus paper in 1821), their taxonomy is still poorly organized. T h e r e are
dozens of named species from all over the world, many of which are known
from partial or fragmentary material, and some of which undoubtedly belong
in different genera. In a 1981 review of the late Jurassic Plesiosauridac, D. S.
Brown recognized four genera and six species: Cryptoclidus eurymems, C. richard50/n', Muraenosaurus leedsii, M. beloclis, Tricleidus seeleyi, and Colymbosaurus trochanteric.
In 1892, H a r r y Seeley first described Cryptoclidus ("hidden clavicle"), named for
the small clavicle bones that rest in shallow depressions on the inner surface
of the front l i m b girdle and are thus hidden from view. Seeley s actual specimen, found in the Oxford Clay* of Bedford, England, has been lost; it
* England's Oxford Clay is one of the most famous and productive fossiliferous regions in
Brown
and
Cruickshank
tgqj).
= surangular; SQ
= premaxilla; PO
= postorhital; PT
= pterygoid; Q
= quadrate;
= squamosal.
Brown
and
Cruickshank
1994).
= surangular; SQ
= premaxilla; PO
= postorhital; PT
= pterygoid; Q
= quadrate;
= squamosal.
The dental
arrangement
of
Cryptoclidus
suggests that this 11-
plesiosaur
lightly built skull, with as many as 100 small sharp teeth that interlocked
items by trapping
them in the
intcrmeshed teeth.
"flying" underwater, with the forelimbs raised on the upstroke and the hind
foot-long
flippers trailing behind: "the flattened body shape provides additional dorsoventral stabilization during subaqueous flight locomotion as in marine
turtles."
In 2002, A r t h u r C r u i c k s h a n k and R. Ewan Fordyce published a detailed
description of a N e w Zealand cryptoclidid plesiosaur that they named Kaiwbekea kaitiki. T h e name, taken from the M a o r i language, means "squid eater of
Katiki." T h e fossilized skeleton was largely complete and included the skull;
all cervical, pectoral, thoracic, and sacral vertebrae; much of the right and left
rib cage; some gastralia; and an almost complete right hind limb and part of
the left. T h e entire specimen was more than 21 feet long, and because the tail
was missing, the animal in life w o u l d have been even longer. T h e authors
wrote that "there is not currently a consensus on plesiosauroid classification;
the diagnoses and content are still debated for the Plesiosauridae, Elasmosauridae, and Cryptoclididae," but based on their cladistic analyses, they
assigned Kaiwbekea to the C r y p t o c l i d i d a e . W i t h its long neck and powerful,
tooth-studded jaws, Kaiwbekea probably fed on fast-moving, m e d i u m - s i z e d to
large prey, probably fishes and cephalopods. T h e structure of the neck vertebrae "provide no evidence of a serpentine mobility, although the craniocervical joint allowed significant movement horizontally and vertically . . . the
degree of ventral movement is hard to assess." ( A s we shall see, the flexibility
of plesiosaur necks is a popular subject for paleontological controversy.) Like
other cryptoclidids (but unlike the elasmosaurs), Kaiwbekea had large eye
sockets, suggesting the ability to hunt deeper in the water column where the
light levels were lower.
W h e n H a r r y Seeley examined a plesiosaur specimen collected from the
Oxford Clay by Charles Leeds of Exeter College, he noted the similarities to
Cryptoclidus but observed that its neck was nearly twice as long. N a m e d for an
eel (Muraena) on account of its long, eel-like neck and for Leeds, Muraenosaurus
leedsii was first described by Seeley in 1874. T h e skull was about 16 inches long,
and the entire animal was estimated at between 15 and 21 feet. Benton (1990a)
wrote that "Muraenosaurus . . . took the long-necked adaptations to an extreme.
S o m e had as many as 70 cervical vertebrae, and the neck could bend around
itself two or three times." W h e r e a s the numerous teeth of Cryptoclidus were
smooth, those of Muraenosaurus were fewer, larger, and vertically ridged. " T h e
hindermost teeth look directly upward," Benton wrote, "but as they approach
the front of the jaw they are necessarily directed more and more outward and
forward." Like Cryptoclidus, its tail was short and its paddles proportionally
small, suggesting that it was a s l o w - s w i m m i n g ambush predator. T h e r e is
another Muraenosaurus species, s u r n a m e d beloclis, referring to the shape of its
clavicle (belemnon is "arrow" or "spear" in Greek; it is also the root of the name
of the cephalopods known as b e l e m n i t e s ) .
T h e Leeds collection also contained disarticulated elements of a plesiosaur that C. W. Andrews (1909) n a m e d Tricleidus seeleyi. Also from the Oxford
Clay, this species is monotypical, meaning that it is the only species in the
genus. It differs from Cryptoclidus and Muraawsaurus in the number of teeth,
which also were more strongly ridged. It too was probably a trap feeder,
dining on soft-bodied cephalopods and crustaceans. Colymbosaurus ("diving
l i z a r d " ) trochanteric, identified and named by Owen in 1840, was the largest
and most massive of the English long-necked plesiosaurs, reaching an overall
length of 22 feet, about a third of which was neck. Because Colymbosaurus is
known only from skeletons without skulls, the skull of Kitnmerosaurus (a new
genus erected by Brown in 1981 and named for the Kimmcridge Clay of
D o r s e t ) m i g h t belong to the same animal, which would make Colymbosaurus a
junior s y n o n y m of Kitnmerosaurus. W i t h M i l n e r and Taylor in 1986, Brown
wrote, " T h e possibility that Kitnmerosaurus m i g h t be s y n o n y m o u s with Coosaurus is discussed, in which case Colymbosaurus would have to be reassigned at family level as a long-necked eryptoelidid." W h a t e v e r its name, this
was another of the U p p e r Jurassic long-ncckcd plesiosaurs that trapped their
prey in a mesh of sharp teeth.
T h e r e are m a n y ways of moving through the water. M o s t marine vertebrates (fishes, sharks, sea snakes, and c r o c o d i l i a n s ) use an oscillating, vertical
tail fin to provide propulsive power. Cetaceans a n d sirenians use an analogous
arrangement, but the movement of their flukes is up and down rather than side
to side. Ichthyosaurs had four flippers, but their caudal fins had developed
into vertical, flattened crescents that moved from side to side and served, like
the tail of a shark, as the propulsive engine. For the four-flippered, short-tailed
plesiosaurs, there were essentially three alternatives. O n e is "paddling," where
the feet of the animal are pulled through the water in the vertical plane; this is
the way ducks swim or the way h u m a n s use their a r m s doing the crawl stroke.
(A nonanimal example is a canoe or, in a form that moves the water without
moving the paddlcr, a watcrwhcel.) In the p a d d l i n g recovery stroke, the duck
closes its webbed foot and brings it forward; the h u m a n s w i m m e r raises his
a r m out of the water. In paddling, the limbs usually alternate the propulsive
strokes, but in principle, they can be used together. " R o w i n g " uses the same
technique of pulling a flattened blade through the water, except that it is in the
horizontal plane and the "oar" has to be "feathered," which means that in the
recovery stroke, it is turned to present its least-resistant face to the water. Few
living animals actually row through the water, but the human breaststroke can
be used as a paradigm. Finally, there is underwater "flying," where the limbs
move at a right angle relative to the direction of motion as the animal moves
forward; this results in a "figure-eight" pattern made by the fin. Examples of
subaqueous "fliers" are sea turtles, penguins, and sea lions. Penguins (like their
terrestrial avian ancestors) use their forelimbs for flying; otariids also use their
forelimbs; and the four-flippered sea turtles use the forelimbs for propulsion
and the hind ones for steering.
T h e locomotion of plesiosaurs has always puzzled and fascinated those
who have studied these extinct marine reptiles. In 1824, Conybeare speculated
that "in its motion, this animal must have resembled the turtles more than any
other," and in 1851, Owen corroborated C o n y b e a r e s opinion. T h e n in 1924,
working with the fossil remains of Plesiosaurus dolicbodeirus, D. M. S. Watson
reconstructed the limb muscles ( i t was the first time anyone had ever done
that with a plesiosaur and decided that they didn't "fly" like turtles but
"rowed." Storrs (1993a) s u m m a r i z e d the myriad discussions of plesiosaur
locomotion and wrote:
T h e model for underwater "flight" for plesiosaur paraxial locomotion is
not a perfect one. Rather, it is probable that the plesiosaurian power stroke
combined elements of "flight" and subaqueous "rowing" with both vertical
(lift-based) and fore-and-aft ( d r a g - b a s e d ) components. Robinson (1975)
was correct in highlighting the hydrofoil-shaped limbs of plesiosaurs.
T h e i r distally tapering configuration created a relatively high aspect ratio
for each limb and this m i n i m i z e d trailing vorticity and lift-induced drag.
Nonetheless sea lions, with limbs constructed in a functionally similar
pattern, do not rely exclusively on lift-induced thrust. . . . An incrtial
gliding phase may have existed between periodic power strokes, again as in
sea lions. At least in pliosaurs, this gliding phase would have been relatively
unencumbered by drag because of their comparatively small surface to
mass ratio (less likely in elasmosaurs whose long necks increased surface
area relative to mass).
W h e n the remains of a large reptile were unearthed by workers at the
London Brick C o m p a n y in Bedfordshire in 1958, work was stopped and
paleontologists from the British M u s e u m were called in. T h e remains consisted of the back part of an animal and the front of the head, the total length
estimated at 36 feet. It was Liopleurodon, and according to Barney N e w m a n and
Beverly Tarlo, the reconstruction enabled them to arrive "at a result different
from other restorations in suggesting a much more streamlined animal."
Although they "discovered evidence that the plesiosaurs had a tail fin on the
upper part of the tail," they d i d not reveal in their 1967 paper what that
evidence was. T h e y did explain, however, why the bones in the " p a d d l e s " of
plesiosaurs are rarely disturbed while the other bones are often scattered:
"they were fixed in cartilage or gristle and were not capable of any movement
on each other, as, for example, our fingers arc." T h e y further concluded that
the forelimbs and hind limbs of pliosaurs functioned differently; the forel i m b s could be raised above the horizontal, but the hind limbs could not,
which indicated to them that the animal could dive in pursuit of food.
In 1975, Jane Ann Robinson wrote an article entitled " T h e Locomotion of
Plesiosaurs," which was filled with sentences such as, " T h e action of this
muscle parallels that of supracoracoid-coracobranchial complex in the pectrum, providing most of the power for adduction." She concluded:
Plesiosaurs, long considered to be rowing animals, are shown to have
employed subaqueous flight, modifying both fore- and hindlimbs into
virtually identical w i n g s . T h e s e can be compared functionally point for
point with the w i n g s of subaqueous flyers (sea turtles, penguins, and
o t a r i i d s ) , but not with the l i m b s of rowing or p a d d l i n g animals. . . .
Differences in l i m b and h e a d / n e c k proportions in plesiosaurs are related
to relative maneuverability; the short-necked forms were agile and pursued
their prey as sea lions do, while the long-necked forms were endurance
s w i m m e r s and not particularly agile.*
* Robinson's paper, published in the Neues Jahrbuchjur Geologic una Palaeontologie in Stuttgart, is
1
Instead of solving the problem, Robinson's paper only opened the floodgates lor a barrage of criticisms and revised interpretations. In a paper published in the same journal in 1982, Tarsitano and R i e s s looked at Robinson's
"elaborate study" and found that her errors had to do with "either incorrect
interpretations of the m o r p h o l o g i c a l data a n d / o r the lack of understanding
of basic biomechanical concepts a n d morphology." In addition, her paper was
"hampered by faulty comparisons with the non-analogous underwater flight
of penguins, sea turtles and sea lions." T h e y wrote that the comparison to the
wings of penguins was flawed because "most tetrapod underwater flyers have
a functional elbow joint, the exception being plesiosaurs and h u m p b a c k
whales."* Tarsitano and R i e s s didn't exactly offer an answer a n d only succeeded in pointing out where R o b i n s o n was wrong. "As yet," they wrote, "it is
unknown how the two sets of wings functioned together in plesiosaurs."
In the same issue in fact, on the next page Frey and R i e s s added a
footnote to the article and wrote, " M u s c l e s inserting at the internal area of
the coracoids or pubices and ischia respectively ( R o b i n s o n 1975) have to be
rejected for anatomical reasons. . . . M o r e likely it seems to us that muscles of
the proximal basis of the limbs m i g r a t e d dorsally to insert at the long neural
spines. Here they could function as levitators. However, this cannot serve as
the answer to the question, why was it necessary for plesiosaurs to develop 4
large 'wings.' All recent tetrapods which fly underwater (i.e., penguins, marine
turtles, sea lions, etc.) are perfectly well-equipped with just 2 wings."
T h e follow-up to Robinson's 1975 article appeared two years later. In the
same Stuttgart journal she published "Intracorporal Force Transmission in
Plesiosaurs," the "second part of a doctoral dissertation prepared at the
University of California at Los Angeles," which might explain the length and
detail of this paper and its predecessor. "Intracorporal force" is the energy, set
up by the motion of the limbs, that ( a c c o r d i n g to R o b i n s o n ) is stored in the
pectrum (pectoral r e g i o n ) , pelvis, and gastralia (belly r i b s ) and helps provide
the propulsive force for the next stroke of the limbs. Robinson compared
* Humpback whales have long flippers, but they use them for steering, not propulsion. Like
all other cetaceans, humpbacks propel themselves with their tail flukes. In a 1 9 9 5 paper, Fish
and Battle discussed the hydrodynamic design of the humpback whale flipper.
that most paleontologists believe that plesiosaurs never left the water (and,
despite the lack of evidence, that they probably gave birth to live young there),
the function of gastralia in the marine reptiles is still unknown. In her
discussion of the locomotion of plesiosaurs, Robinson (1977) suggested that
the gastralia functioned as a component of a dorsoventral stabilizing system
(the "archers bow construction") that absorbed the up-and-down m o t i o n of
the fore and hind limb girdles while transmitting the propulsive force to the
body. However, this complex explanation has been questioned, most recently
by L i n g h a m - S o l i a r (2000c), because of the ineffectiveness of an inflexible
backbone in storing strain energy in something like an archer's bow. Instead,
he suggests that the ribs (and g a s t r a l i a ) aided force transmission more like the
cables in a suspension or bowstring bridge from fixed supports analogous to
the plesiosaur backbone.
In a letter written to me in January 2002, L i n g h a m - S o l i a r articulated his
ideas about the function of gastralia:
In early ichthyosaurs, especially massive-bodied forms, robust gastralia
probably provided support, whereas in the later streamlined or thunniform
ichthyosaurs, lighter more delicate gastralia m a y have played a very i m portant part in maintaining an advanced hydrodynamic body shape. In
plesiosaurs, where large ventral body mass ( g u t ) was concentrated over a
relatively short body length the gastralia may have played a vital role in
s u p p o r t . . . . Furthermore, a more rigid thorax is essential in a paraxial flyer.
Gastralia are probably made redundant in birds by the enormous sternum
and in marine turtles by the plastron. In the latter group a rigid plastron
(the lungs are dorsally placed a n y w a y ) almost certainly meant that breathing was facilitated by the enormous pectoral musculature during contraction and expansion (possibly a good analogue for plesiosaurs). Mosasaurs,
in contrast, lacked gastralia. Being more slender and elongated the ribs
were probably adequate to assist in breathing and in maintaining body
shape. Furthermore the ventral mass was probably better spread over the
length of the animal. In enormous mosasaurs such as Mosasaurus hoffmanni
the thoracic and anterior abdominal ribs are extremely deep, offering a
good deal of support. T h e disadvantages of gastralia in an advanced
aspect ratio
10
10
units
wide has an aspect ratio of one, whereas one that is 20 units long and 5 wide has an aspect
ratio of four. One of the reasons inclined planes (and wings) with high aspect ratios have a
higher lift-to-drag ratio is that they generate less of a vortex at the tips. The lift-to-drag
ratio of a plane is also increased by its having a streamlined profile . . . a streamlined body
experiences lower drag forces than one with a rectangular profile.
conclude that these creatures were unable to dive after their prey. Furthermore, we can see from the structure of the shoulder girdle that the associated muscles that drew the limb forward were just as strong as those which
drew it back. T h i s means that these long-necked plesiosaurs could twist
and turn extremely well by combining a normal swimming stroke of one
limb with a backing stroke of the other. T h i s sort of action was simply not
possible for a pliosaurit did not have sufficient muscle power for an
effective backing stroke.
In a 1994 study of the s w i m m i n g capabilities of the marine reptiles, M a s sare wrote that "the two lineages of plesiosaurs, the plesiosauroids (Superfamily Plcsiosauroidca) and the pliosauroids (Supcrfamily Pliosauroidca)
shared a similar mode of locomotion, that of subaqueous flying, using their
two pairs of winglikc appendages as hydrofoils," but the swimming of plesiosaurs is not that simple. In 1975, Robinson argued that the animals "flew" like
overgrown penguins but where penguins have only two flippers, the marine
reptiles had four, which presented a more complicated problem in hydrod y n a m i c s . According to Massarc's 1994 summary:
T h e old idea of subaqueous rowing assumed that the two pairs of wings
would move simultaneously. W o u l d underwater flyers with an asymmetric
power stroke necessarily beat their four limbs in unison? Ricss and Frcy
argued that the two pairs of wings were out of phase by half a stroke, such
that the forelimbs were in the power phase (backward, downward movem e n t ) while the h i n d l i m b s were in the recovery phase (forward, upward),
and vice versa. T h u s the plesiosaur was always generating thrust. Alexander
pointed out, however, that swimming would be very inefficient with the
fore and hind limbs out of phase because the h i n d l i m b s would be accelerating water that was already in motion. It would be more efficient for the
fore and hind limbs to work together and simultaneously accelerate a larger
mass of water.
Bakker (1993a) wrote that "the plesiosaur body was short, compact, oval in
cross-section, and strongly-braced below by thick central ribs that resisted the
compressional forces generated by the pulsating cycles of the swimming
response to
niv
atheoretical i n
his
Recent workers have regarded the four wings of plesiosaurs as four organs
performing essentially the same function, i.e., an integration of the components of thrust, lift, stability, and steering in one and the same system. In
extant vertebrates on the other hand (e.g., sharks, whales, dolphins, ichthyosaurs, flying fish, etc.) and in machines (airplanes, powered gliders) the
efficiency of lift and thrust components is m a x i m i z e d by largely independent g e n e r a t i o n . . . . In birds the same organ performing distinctly different
functions accounts for the complexities of their wing structure. In a number of large marine vertebrates e.g., lamnid sharks, whales, dolphins, and
laivc ichthyosaurs, efficient locomotion is dependant upon a separate pair
of h y d r o d y n a m i c organs to achieve stability, passive lift, and steering.
L i n g h a m - S o l i a r dismisses Robinson's elaborate "archer's bow theory,"
whereby an inflexible bow composed of the dorsal and ventral elements of the
skeleton serve as the basis for propulsive forces on the four hydrofoils during
underwater flight, and also Riess and Frey's (1982) theory of passive upstroke
of the limbs. He replaces them with his own synthesis, published in great
detail in Neues Jahrbuch and then as a popular article in the British newspaper
The Guardian on November 16, 2000:
Traditionally all four plesiosaur limbs have been treated as identical structures. T h i s is far from so. T h e anterior limbs are swept back as in swallow's
wings the posterior ones are relatively straighten Research has shown that
the swept-back or crescent shape is d y n a m i c a l l y more efficient than the
straight wing for flight. T h e r e is no elbow or wrist-joint in plesiosaur
limbs, so the ability to make delicate changes in pitch, direction etc., was
limited. Of further significance is the way in which marine animals such as
sharks and ichthyosaurs rise or descend in the water by elevating or lowering the leading edge of the paddles. However, lift in a hydrofoil works best
theoretically weak, so was Riess and Frey's 'passive supination' hypothesis, the gastrolith and
ballasting hypothesis is highly dubious . . . etc. Most later theories were simply taking earlier
statements of four-wing (light as read (undoubtedly an alluring proposition), despite the
fact that it was never really soundly investigated or tested in the light of alternative
theories in truth they depended on a wing and a prayer (pun intended)."
when it is inclined at a precise angle to the water How known as the angle
of attack. Hence, the wing's capacity to act simultaneously as a rudder and
flight organ is impractical. For a pursuit predator chasing elusive prey, this
would be disastrous. T h e solution? Sharing flight functions between the
anterior and posterior limbs. T h i n k of plesiosaurs as using a front-wheeled
drive engine (thrust and lift) with the steering and a n u m b e r of other
functions at the back (ideal for rotating and maneuvering because of their
greater distance from the centre of b a l a n c e ) . . . . At other times the hind
limbs could also be used in rowing the predominantly ventral m u s c u l a ture would allow this. W h i l e the g l a m o r o u s notion of plesiosaurs as four
rather than two-winged fliers is appealing, reality suggests a very effective
division of functions between the anterior and posterior limbs, necessitated by mechanical and anatomical limitations, but every bit as unique.
In a book revcalingly called In the Presence of Dinosaurs, John C o l a g r a n d c
resurrects the lifestyle of various long-extinct reptiles, with only a brief nod to
the idea that he might be m a k i n g it all up. In the introduction, he says, " T h e
behaviors we ascribe to the a n i m a l s are by nature speculative, but no more
'radical' than some of the forms of behavior exhibited by m o d e r n animals." In
other words, because some living animals exhibit enigmatic behavior, there is
no problem in fabricating equally strange behavior for creatures that have
been extinct for 100 million years and arc known only from fossils, some of
them mere fragments. Here is Colagrande's account of the egg-laying procedure of clasmosaurs:
T h e y normally start approaching just before sunrise, while the m o o n is
still above the horizon. Like an invading armada, scores of the giant
reptiles head for the beach, enter the surf zone, thrust themselves out of
the foam, and laboriously pull themselves onto the sand with a loud
grating hiss. Every once in a while one of the leviathans will stop m o m e n tarily and with a shake of its head, snort a great spray of salty water from
its nostrils. S o m e marine reptiles absorb great amounts of salt, which can
kill them if allowed to build up. Special glands in their heads remove the
excess salt from their blood and d u m p it into their nasal passages, where it
is unceremoniously expelled.
their prey, which consisted of other swift marine reptiles such as ichthyosaurs,
as well as large fishes and perhaps even cephalopods. Did pliosaurs row or fly
through the water? W a s their s w i m m i n g similar to that of turtles? Sea lions?
Penguins? Did they pull with their forelimbs and let their hind limbs trail
behind, or did they flap hind and forelimbs in some sort of synchrony? Did
the long-necked plesiosaurs locomote differently from the short-necked
pliosaurs? Did any of them ever come out of the water?
H a r r y Seeley, who in 1880 was one of the first to recognize fossilized unborn ichthyosaurs, later claimed to have discovered the same for plesiosaurs.
In 1888, he published a short note in which he described "four more or less
complete specimens regarded as foetal plesiosaurs, together with fragments of
at least three others. T h e y are remarkable for having the flesh mineralized with
phosphate of lime, and still show many of the characters of the external form
of the body, but slightly distorted by decomposition. O n l y one individual has
the head preserved; its extreme length is about 14 mm [Vz i n c h ] , . . . Hence, the
author regarded this specimen as showing that Plesiosaurus was viviparous,
and that in one species from the Lias many were produced at one birth." It is
likely that plesiosaurs were viviparous, but Seeley s "evidence" cannot be used
to demonstrate it. W h e n R. A. T h u l b o r n (1982) decided to reexamine the
material in London's Natural H i s t o r y M u s e u m that Seeley had described, he
found that it consisted of "an irregular nodule of pyritic mudstone and shale;
a small fragment, broken from the main nodule; five slides each with a thin
section; a slide with twenty-three stained sections of a modern lizard embryo."
Examining the material that Seeley had so carefully identified as heads, necks,
bodies, tails, and limbs, T h u l b o r n found "no evidence of organic structure in
any of the supposed embryos . . . and some of the anatomical features
identified by Seeley are no more than surface irregularities, scratches, pits, and
similar weathering effects."
W h a t , then, were Seeley's "embryos"? T h u l b o r n reinterpreted the original
fossils as a crustacean burrow system, with "only one structural feature (the
'dorsal ridge' noted by S e e l e y ) that I have been unable to match in any other
thalassinoid burrows." In his 1994 discussion, Storrs wrote, " T h e so-called
'embryos' had been acquired by Seeley after passing through the hands of a
local fossil dealer. T h e y had been, furthermore, highlighted or 'enhanced' by
vertical tail fins.) A n d if we can't tell how an aquatic reptile moved in the
water, we are utterly in the dark about how or even ifthese great reptiles
moved on land. For example, Michael Taylor wrote, "Plesiosaurs may have
given birth to live young or laid eggs in pits scraped out with their tails." N o t e
the use of the words "may have," which indicates that Taylor is only guessing
about what the plesiosaurs might have done a far cry from Colagrande's
descriptive phrasing, which makes it sound like he has actually observed
plesiosaurs laying eggs or hunting fish. S o m e paleontologists delight in speculation about the way their subjects walked or swam or flew; it gives life to the
otherwise mute fossils. But to describe in unequivocal terms how a longextinct animal like a plesiosaur caught its prey does a disservice to the science
of p a l e o n t o l o g y by removing the element of mystery and replacing it with an
ill-conceived and unwarranted certainty. W i t h o u t a time machine, we can
only wish that we knew how they hunted.
T h e long-necked plesiosaurs were probably among the slowest of the
ancient marine reptiles, with their barrel-shaped bodies creating substantial
drag. Judy Massarc, in her 1988 analysis of the s w i m m i n g capabilities of
M e s o z o i c marine reptiles, wrote that "the plesiosauroids . . . do not fall within
the range of o p t i m u m fineness ratios. T h i s would have resulted in greater drag
for a given size than for ichthyosaurs and pliosaurids, and suggests that the
more elongate forms probably had slower continuous s w i m m i n g speeds."
Lateral movements of the long neck would also affect the plesiosaurs' ability
to steer, creating considerable yaw (deviation from a straight course) as the
animals plowed through the water. M o v e m e n t of the head and neck as the
animal paddled must have been the functional equivalent of having a rudder
on the front end, requiring constant course corrections. It is possible that the
animals compensated for these deviations by developing a s w i m m i n g style
that was constantly veering from one heading to another, because it seems
certain that they could not have moved in a straight line if the head was in
motion from side to side or up and down. W i t h a long, snaky neck and a
mouthful of sharp teeth, the "swan lizards" (Bakker's 1993 name for t h e m )
were almost certainly fish eaters, but how they captured their prey is not
immediately evident. D i d they take a breath and dive after fishes (the way
some toothed whales do t o d a y ) , or did they float at the surficc with head and
neck hanging down underwater, like a snake hanging from a branch? Did they
float at the surface with the head raised, and then plunge it downward to
capture a fish in the manner of herons and egrets?*
S o m e have assumed that plesiosaurs, because of their incredibly long
necks, could coil their heads back and strike at passing prey like a snake. In
1897, Charles R. Knight illustrated Elasmosaurus platyurus with its neck curved
into a figure eight, looking like a python grasping its prey.f "But," argued
Samuel W i l l i s t o n (1914), "the plesiosaurs could not and did not use their neck
in such ways. T h e y swam with the necks and head, however long, directed in
front, and freedom of movement was restricted almost wholly to the anterior
part. T h e posterior part of the neck was thick and heavy, and could not have
been moved upward or downward to any considerable extent and not very
much laterally." (In a letter to me in 2002, Ken Carpenter wrote, " W h e n the
Elasmosaurus mounts were done at Denver, I played with the neck to determine
the true range of motion, not with sketches on paper. I concluded that
W i l l i s t o n was right.") M i c h a e l Benton, author of The Reign of the Reptiles (1990)
and Vertebrate Paleontology (1997), the latter a respected textbook, disagrees as
do many other paleontologists. In Reign of the Reptiles, he wrote ( o f elasmosaurs
of the Jurassic, such as Muraenosaurus), " S o m e had as many as 70 cervical
" The stud)' of fishing birds actually oilers little help in determining how plesiosaurs might
have ted. Swans, with comparably long necks, are nor fish eaters; they churn up aquatic
plants from the bottom. Flamingos, with even longer necks, stand on their stiltlikc legs and,
with their heads upside down, filter tmv organisms from the water with a sievclikc arrangement in their beaks. Cormorants and anhingas (the latter sometimes known as "snakebirds" provide perhaps the best analogue; they dive beneath the surface and swim alter fish,
capturing them with their beaks. The cormorant grabs its prey, but the anhinga spears it
and then, rising to the surface, (lips it up and catches it headfirst. The long-necked herons
and egrets are not swimmers at all but stand in the shallows, poised to snatch swimming fish
or frogs.
t Knight's illustration was made under the direction of E. D. Cope for the American
Museum of Natural History In their 1 9 8 2 book about Knight (Dinosaurs, Mammoths and
Cavemen), Sylvia Czerkas and Donald Glut wrote, "The depiction of the extremely serpentine neck, although now known to be an error, was based on the evidence available to Cope
at the time. Despite this inaccuracy, the painting is a stunning work of art conveying the
drama of Mesozoic marine life."
vertebrae, and the neck could bend around upon itself two or three times.
T h e elasmosaurs no doubt jabbed their snake-like necks rapidly among the
scattering clouds of teleost fishes. T h e y could have darted the head in and out
and seized several fish without moving the body at all." It is known that they
swallowed stones for ballast (or to help grind their food), so they might have
sunk to the bottom to wait for unwary fish to come into striking range. T h i s
scenario is supported by the position of the eyes in the skull, facing slightly
upward.
In contrast, N e w m a n and Tarlo (1967) wrote, "in the long-necked plesiosaurs, both sets of limbs could only move in a horizontal plane, and since they
could not raise their forelimbs, they could not dive." T h e y concluded that
"the long-necked forms were adept at twisting and turning with great speed,
but all their time was spent at the surface of the sea." In his 1968 book The
Pattern of Vertebrate Evolution, Halstead (a.k.a. Tarlo) wrote, " T h e inevitable
conclusion is that the head and greater part of the neck were lifted out of the
water during rapid manoeuvring, swinging over the surface and dropping in
again on the sighting of fish. T h i s high degree of manoeuvrability not only
enabled the long-necked plesiosaurs to fish efficiently but must also have been
equally advantageous in helping them to escape the attention of predators."
As with their method of locomotion, the plesiosaurs have managed to turn
the simple business of feeding into another palcontological conundrum.
A subject not often mentioned with regard to the elasmosaurs is the great
degree of vulnerability represented by the long, exposed neck. Very long necks
are rarely evolved because they are costly in terms of growth and maintenance,
as well as the extra respiratory work needed to breathe through elongated
trachea. Because all the energy spent on a longer neck means less energy that
could go elsewhere tail, limbs, growth, parturition, tougher hide, and so
on there must have been a significant metabolic advantage to a long neck. It
might have enabled the animal to catch enough food to support this anomalous structure, but this seems like circular reasoning: an animal would not
have developed a long neck so it could harvest the energy necessary to
support such a structure. Moreover, the neck is one of the most vulnerable
spots in an animal, since the nervous, circulatory, and respiratory systems all
run through it in close proximity. W i t h such dangerous predators as giant
The long-ncckcd tcrrcstn.il dinosaurs faced the same problems. We do not know how (or
even i f ) creatures such as Apatosaurus, Rrachiosaurus, and Diplodocus raised their heads to such
astounding heights, because studies of the biomechanics of their skeletons have so far
proved inconclusive. With the carnivorous thcropods such as Tyranncsaums rex and Allosaurus
on the prowl, they too would have been extraordinarily susceptible to attack.
me to state that there is not a shadow of a doubt that the plesiosaurs, both
Cretaceous and Jurassic, had the habit of swallowing such stones. At least
thirty instances arc now known of the occurrence of the very peculiarly worn
pebbles between the ribs or with the remains of plesiosaurs in Europe and
America. T h e fact was first published by Professor Seeley in England, in
>8 ."
77
than vision in air because they had to capture their prey underwater and had
to see above the surface only when they breathed or rested. Tbalassiodracon is
known only from the late Triassic to early Jurassic shales of England.
Some plesiosaur fossils, alter lasting lor 100 million years, did not survive
the blitzkrieg bombing ol England during World War 11. Plesiosaurus conxbean
was first described in 1881 from C h a r m o u t h , England, and kept in the collection of the Bristol C i t y M u s e u m . It was an almost complete skeleton, lacking
only the tip of the tail, and remarkably, what appeared to be an impression of
the skin was preserved in a thin film over the body. T h e r e were no scales or
scutes, so this species and perhaps the other plesiosaurs may have had
smooth skin, in contrast to most living reptiles, which have scales of one sort
or another. T h e fossil was destroyed in 1940, but the British M u s e u m ( N a t u ral H i s t o r y ) in London had made a cast, and it survived the raids. Bakker
(1993a) examined it and reclassified Plesiosaurus conybeari as Attenborosaurus (after
filmmaker David A t t c n b o r o u g h ) , because "it is the only plesiosaurian of any
age that combines a very long neck with a long, massive, evenly tapered snout,
and very large, conical tooth crowns."
In their 1999 description of Plesiosaurus tournemirensis ( n a m e d for the southern French village of T o u r n c m i r e ) , Bardet, Godefroit, and Sciau wrote:
During the M i d and Late Jurassic, elasmosaurs apparently began to spread
all over the world, as they have been reported from Argentina, W y o m i n g ,
and India. . . . During the Late Cretaceous, elasmosaurs achieved a worldwide distribution as they have been found in both the northern and
southern hemisphere. T h e y are especially abundant in N o r t h America,
where most of the genera have been described, but they also occur in
western Europe, Russia, Japan, Africa and the M i d d l e East, M a d a g a s c a r ,
South America, N e w Zealand, and Antarctica. T h e Elasmosauridac d i s a p peared at the end of the Maastrichtian, during the K / T biological crisis.
In a 1994 study of the extinction of the marine reptiles, N a t h a l i e Bardet
noted that by the mid to late Triassic (234.3
to
zz
placodonts and the mixosaurid ichthyosaurs were gone; by the late Jurassic
(150.7 to 144.2 million years a g o ) , the cryptoclidid plesiosaurs and the family
Ichthyosaundae were gone; and in what is known as the early late Cretaceous
One
century
nineteenthdescription
of a plesiosaur
was
Plesiosaurus
conybeari, whose
(98.9 to 93.5 million years a g o ) , the last of the ichthyosaurs (the Platyp t e r y g i d a e ) were gone. Sixty-five million years ago (the end of the Cretaceous), when the C h i c x u l u b asteroid hit, the pliosaurs, elasmosaurs, and
mosasaurs, along with the terrestrial dinosaurs, disappeared forever. A high
incidence of extinction among marine invertebrates (ammonites, bivalves,
corals) has been detected at the end of the Jurassic (144.2 million years ago),
but it is not known whether these extinctions contributed to the downfall of
the placodonts and mixosaurs or whether the same thing that eliminated the
invertebrates also wiped out the reptiles. W h e n the C e n o m a n i a n stage of the
Cretaceous period ended 93.5 million years ago, there was another mass
extinction of marine invertebrates ( R a u p and Scpkoski 1986), but again, the
cause-and-effect relationship cannot be established. Bardet said, " T h e fact
that plesiosaurs may have been more opportunistic predators than ichthyosaurs could explain vvhv thev have not been directly affected," but whatever the reason, the ichthyosaurs died out long before the plesiosaurs did. T h e
asteroid that struck the earth 65 million years ago was somehow responsible
for a precipitous drop in the phytoplankton of the oceans' surface, and this
may have caused a massive break in the food chain, the effect of which was felt
all the way up to the top predators, the plesiosaurs and mosasaurs.
In a 1993 article called "Plesiosaur Extinction Cycles," Robert Bakker wrote,
"It is well known that the terminal Cretaceous event exterminated all plesiosaurs and all mosasaurs in the Western Interior [ S e a w a y ] , as well as elsewhere.
However, most traditional plesiosaur classifications give the impression that
this mass extinction was a most unusual event and that, for most of the
Jurassic and the Cretaceous, the major plcsiosaurian clades had been evolving
continuously and without serious disruption." T h i s , Bakker believes, was not
at all the case; instead, "plesiosaur history was punctuated by a series of
sudden mass extinctions that define natural units of marine history." He lists
successive radiations and extinctions, beginning with the "Terminal Tnassic
Extinction and Early Liassic R e - R a d i a t i o n " and concluding with the " M i d Cretaceous Extinction and R e - R a d i a t i o n , " which eliminated the ichthyosaurs
and opened new opportunities for plesiosaurs. Extinctions of some dinosaur
species on land occurred at the same time the Jurassic-Cretaceous boundaryand Bakker points out that roughly in the m i d d l e of the period, there was
"a marked turnover among herbivorous dinosaurs the Iguanodontidae are
replaced nearly entirely by the Hadrosauridae. . . . All the available evidence
favors the view that the replacement at the family level among dinosaurs
occurs at the same general interval as do the extinction events among large,
specialized marine reptiles." As to the identification of the extinction agent, he
wrote, "Draining of the cratonic interior and continental margins would
remove much of the productive area of the shallow sea." In other words, when
the Western Interior Seaway dried up, the animals that lived in it died off.
In their discussion of plesiosaur remains found in the M o n s Basin of
southern Belgium, M u l d e r et al. (2000) also summarize the ( l i m i t e d ) material
on Maastrichtian elasmosaurs worldwide. R a r e in the M o n s Basin region,
they are much more c o m m o n in Cahfornian and M o r o c c a n deposits of the
same age and type, that is, late Cretaceous marginal seas such as the coast of
California, the Western Interior Seaway, the Mississippi Embayment, and the
N o r t h Atlantic Basin. T h e authors note that Bakkcr (1993b) suggested that
"the absence of fast-swimming plesiosaurs could be explained by the presence
of an exceptionally dense algal forest. Only the California coast had an abrupt
continental slope, with a narrower zone of shallow water where an algal forest
could grow." M u l d e r and his colleagues, however, believe "that the presence of
an abrupt continental slope also coincided with an upwelling of nutrientenriched water, which favored the presence of a high biomass, being an ideal
condition for open-water predators such as clasmosaurids." T h e worldwide
distribution of elasmosaurs indicates that, like the mosasaurs, they were
widespread and diversified during the late Maastrichtian, the end of which (65
million years a g o ) was marked by an explosion of an extraterrestrial object
that hit in what is now the Yucatan. T h e y conclude, therefore, that "the
extinction of the elasmosaurids at the K / T boundary thus appears to be
sudden rather than gradual."
The Pliosaurs
One of the relatives of the Plesiosaurs, the Pliosaur, of which genus several species of great size
are known, perhaps realized in the highest degree possible the idea of a huge marine predacious
reptile. The head in some of the species was eight feet in length, armed with conical teeth afoot
long. The neck was not only long, but massive and powerful, the paddles, four in number, were
six or seven feet in length and must have urged the vast bulk of the animal, perhaps forty feet in
extent, through the water with prodigious speed.
powerful heart and lungs. Imagine such a creature raising its huge head twelve feet or more out
of the water, and rushing after its prey, impelled with perhaps the most powerful oars possessed
by any animal. We may be thankful that such monsters, more terrible than even the fabled seaserpent, are unknown in our days. J. W. Dawson (1903)
T h e r e arc those who would with good reason combine the plesiosaurs
with the pliosaurs, but there are enough differences in functional morphology
to separate them, at least for this nontechnical discussion. Appearing first in
the early Jurassic, the short-necked pliosaurs developed into the apex predators of the M e s o z o i c , gobbling up sharks, large squids, ichthyosaurs, plesiosaurs, and probably smaller pliosaurs as well. W h e r e a s the teeth of plesiosaurs
were often narrow and needle-like, as befits fish eaters, those of the shortnecked pliosaurs were conical, massive, sharp, and ridged. T h e i r heads were
often disproportionately large, and their jaws were powered by huge muscles.
T h e name pliosaurs means "more lizardlike" and was bestowed on them by
R i c h a r d Owen in 1842, because they appeared to h i m to be even more lizardlike than the plesiosaurs. He wrote, " T h e Enaliosaurs* are immediately connected with the Crocodilian reptiles by an extinct genus, represented by
species of gigantic size. T h e Reptile in question is essentially a modified
Plesiosaurus, but its modifications appear to entitle it to be regarded as a
distinct genus, which, as it is more closely allied to the true Sauria, I propose
to call it Pliosaurus."
One reason to combine the plesiosaurs with the pliosaurs is the existence
of some forms that seem to be intermediate between the two, such as Macroplata ("long blades," named for its scapulae), which was either a short-necked
plesiosaur or a (relatively) long-necked pliosaur. Based on its powerful forelimbs (attached to large bony plates that braced the front paddles), it was
probably a fast swimmer. It had an elongated, crocodile-like skull that was 28
inches long and a tapering, rather inflexible neck composed of 29 short,
flattened cervical vertebrae. Its total length was about 16 feet. Tate and Blake
(1876) originally named the first recognized species Plesiosaurus longirostris, but
the name Macroplata longirostris ("long r o s t r u m " ) was applied by W h i t e in 1940.
T h e second species of similar size (15 feet total l e n g t h ) was Macroplata tenuiceps
("narrow h e a d " ) from the Lower Lias ( H e t t a n g i a n ) of Warwickshire, En* Enaliosaurus
enalios,
and was introduced hy Owen in i8;q as a catchall term to include all the ichthyosaurs and
plesiosaurs known at that time. "As we shall sec later," wrote Williston ( 1 9 1 4 ) , "the
plesiosaurs are really of remote kinship to the ichthyosaurs, and there is no such natural
group as the Enaliosauria."
Mr.
and
Mrs.
Macroplata
tenuiceps
looking
in
thejossil
record to indicate
gland. Like M. longirostris, it had a neck about twice the length of its sharp-
females in this
had a shoulder girdle with coracoids that were proportionately much larger
than the scapulae, indicating a powerful forward stroke for fast swimming.
certainly
As Darren Naish wrote in a letter to me, " M o s t workers now agree that there
Both sexes
about
but this does not mean goodbye to the Plesiosauroidea and Pliosauroidea.
S o m e plesiosauroids are short-necked, and likewise some pliosauroids are
long-necked."
Another possibly "intermediate" form was originally named Plesiosaurus
arcuatus by Owen in 1840, but an examination of the material, particularly the
skull, convinced Arthur Cruickshank (1994b) that it should be reclassified
Eurycleidus arcuatus (arcuatus means "curved" and refers to the shape of the
vertebrae), as per the 1922 analysis of Andrews. T h e 13-foot-Iong specimen,
which probably came from the Lyme R e g i s area, "seems to possess a suite of
characters intermediate between those of the pliosauroids and the plesiosauroids . . . [which is] important and interesting in itself, as further study of
possible.
ij feet
were
long.
more complete Lower Jurassic plesiosaurs might help to elucidate the process
of change from one superfamily to another." T h e back of the skull of Eurycleidus resembled that of the plesiosauroids, but the large teeth in the front
were indicative of pliosaurs. Cruickshank wrote, "Because of the lack of
fusion of significant skull elements [and other anatomical differences], it is
confirmed that this is not only a small specimen of the species Eurycleidus
arcuatus, but most likely an immature juvenile." In his 1996 review of the shortnecked plesiosaurs of N o r t h America, Kenneth Carpenter wrote: "Traditionally, all short-necked plesiosaurs have been grouped together into the
Pliosauridae. However . . . the term 'pliosaur' should not be used indiscriminately for any short-necked plesiosaur. . . . I therefore conclude that the
short neck has appeared independently at least twice in the Plesiosauria, and
the t e r m 'pliosaur' to refer to any short-necked plesiosaur should be abandoned to avoid any phyletic implications."
Archaeonectrus rostratus, previously known as Plesiosaurus rostratus, was a very
primitive pliosaur with a wide geographic range, with fossil remains from
the Lower Lias ( S i n e m u r i a n ) of England and possibly the early Jurassic of S i beria. T h i s 12-foot-long animal resembled the true pliosauroids in its large,
elongated head, and for an early Jurassic form, it had an unusually short neck,
only about one and a half times the length of the skull. Its teeth were pointed,
conical, and curved back. In typical pliosaurid fashion, the forelimbs were
somewhat smaller than the hind ones. S o m e of the centra of the tail vertebrae
were vertically compressed, suggesting a vertical tail fin. It is not clear whether
Archaeonectrus represents the earliest member of the Pliosauridae or if it is
simply an early type that independently evolved pliosaurid-like features. In his
1999 Plesiosauria Translation and Pronunciation Guide, Ben Creisler summarized the
plesiosaur-pliosaur debate:
Traditionally, the small, short-necked, large-headed plesiosaurs have been
considered "pliosaurs," and classified either in the family Pliosauridae itself
or as a distinct family ( P o l y c o t y l i d a c ) in the superfamily Pliosauroidea.
N e w research questions this assumption. Carpenter (1996, 1997) interprets
polycotylids as a group of short-neck plesiosauroids, closely related to
elasmosaurids based on c o m m o n features in the skull; Bardet (1998) has
a complete skull of
P.
brachyspondylus
in
1980
From
an
analysis of
paleontologists
20-foot-long
and reptiles with deep bites with its strong caniniform anterior dentition, and
Rhomalcosaurus
then used its broad, unobstructed palate, widened posterior gape and hooked
posterior dentition to help move the prey down the gullet." In other words,
gative
today. It is thought
that other plesiosaurs
relied primarily
sight.
on
anything and everything that it encountered including dead dinosaurs. During the excavation, three armored scutes (horn-covered bony plates set in the
s k i n ) were found, obviously not from a pliosaur. Taylor, N o r m a n , and
C r u i c k s h a n k (1993) identified them as belonging to an unidentified armored
ornithischian dinosaur such as an ankylosaur or a stegosaur and tentatively
concluded "that the pliosaur had been scavenging a dinosaur corpse shortly
before its own death, and that the scutes were transported inside the p l i o s a u r s
stomach."
One of the more spectacular of the English pliosaurs was Rbomaleosaurus
(from the Greek rhomaleos, meaning " s t r o n g " ) . T h e first Rbomaleosaurus fossil
was found in an alum q u a r r y in Yorkshire in 1848 and described by Alexander
Carte and W. H. Baily in 1863. ( T h e y named it Plesiosaurus cramptoni, but it was
renamed Rbomaleosaurus zetlandicus in 1874.) T h e magnificent specimen, 23 feet
in total length and complete except for one of the flippers, which had been
destroyed before the fossil was spotted, was displayed for five years in M u l grave Castle, home of the M a r q u i s of N o r m a n b y , owner of the alum mine. In
1853, the fossil was brought to Dublin, where it was displayed at the Z o o l o g i cal Society of Ireland, whose annual report included this description:
It was presented by the M a r q u e s s of N o r m a n b y to S i r P h i l i p C r a m p t o n ,
who has kindly put it at the disposal of the Society for exhibition. T h e
most interesting relic, one of "the great Sea Dragons" of the ancient world,
will no doubt, prove eminently attractive, not only to the citizens of
Dublin, but to the many scientific and other visitors likely to visit this city
during the next few months. Its size is so great that the C o u n c i l felt obliged
to construct a special budding of 36 feet long for its due exhibition.
After its original triumphs, Rbomaleosaurus was exhibited in various buildings of what is now the N a t u r a l H i s t o r y M u s e u m of Ireland and finally
broken up. "For those who want to see the specimen," wrote Roger Osborne
in his idiosyncratic discussion of Yorkshire's geological and paleontological
history, "there is bad news and good news. T h e bad news is that the body of
the fossil is broken up and kept in crates in storage
during the last 140 years several casts of the specimen have been taken, and
these can now be seen in the Natural H i s t o r y M u s e u m in London, the Bath
Literary and Scientific Institution, and Cornell University in N e w York."
W h a t sort of animal was this "great sea dragon"? It was one of the earliest
of the giant sauropterygian predators that "fed on a w i d e variety of active
prey, and forcibly dismembered larger prey by shaking and twisting them"
(Taylor 1992). For some 30 or 40 million years, the 18-foot-long, crocodile-
headed rhomalcosaurs were among the top predators in the ocean. T h e i r jaws
ended in a flattened, spoon-shaped structure ( s o m e t i m e s known as a rosette)
a r m e d with particularly large tusks, probably useful in getting a grip on the
prey before twisting off a piece. As expected in aquatic animals, Rhomaleosaurus
had no eardrum. T h e t y m p a n u m is an adaptation to hearing airborne sound,
and a primarily aquatic animal has less need for a t y m p a n u m because it can
hear by direct conduction from the water. " T h e ears were not acoustically
isolated from the braincase," wrote Taylor (1992), "so underwater directional
hearing was poor, and sonar was not possible." T h e large eyes suggest that
they were visual hunters, and the presence of a sclerotic ring meant that
they could adjust the shape of the eyeball underwater for increased visual acuity. Rhomaleosaurus zetlandicus and perhaps the short-necked Cretaceous polycotylids and the elasmosaurs had additional modifications that made them
even more efficient predators. In his detailed analysis of the head of R.
zetlandicus, Taylor (1992) concluded that it could swim with its mouth slightly
open, allowing water to pass through scoop-shaped openings in the roof of
the mouth, then through a channel where smell sensors were located, and out
through the external nostrils. T h i s arrangement would have enabled this 18foot plesiosaur to hunt its prey by smell like a shark ( C r u i c k s h a n k et al.
1991). Brown and C r u i c k s h a n k (1994) suggested that Cryptoclidus also breathed
through its mouth, and like all rhomalcosaurs, it could detect scents underwater. But unlike the powerful pliosaurid Rhomaleosaurus, the skull and jaws of
Cryptoclidus were lightly built, and it probably fed on soft-bodied cephalopods
and small fishes. Brown and C r u i c k s h a n k concluded, " T h e general structure is
compatible with a filter feeding habit suggested for cryptoclidids by Brown
(1981a) and resembles that of modern 'krill' feeders described by Massare
(.987)."
One of the most unusual plesiosaur fossils in fact, one of the most
unusual fossils of anything is the plesiosaur found in 1986 in an underground
opal mine at C o o b e r Pedy, South Australia. T h e fossil, which was in thousands of pieces, was found by a miner named Joe V i d a , whose clumsy attempts to excavate it resulted in many of the smaller pieces being lost. Paul
W i l l i s painstakingly reassembled it, and it is now an almost complete skeleton
of a small plesiosaur about 6 feet long, with the skull, lower jaw, teeth,
vertebral column, ribs, and most of the pelvic and pectoral girdles composed
"Eric,"
Australian
called "Eric the Half-a-Bcc." ( T h e opalizcd fish gastroliths and vertebrae that
plesiosaur
opalized
dish bought "Eric" and paid W i l l i s to restore it, but he went bankrupt and
put the opalizcd fossil up for public auction. Because the entire fossil was
made of precious opal, its value was estimated at $300,000, and there was no
way of guaranteeing that someone would not buy it and cut it into small
pieces. ( O p a l is a mineral that consists of silica that has filled-in fissures and
cavities of rock. It is usually colorless, but in g e m opal, tiny silicate m i c r o spheres reflect light in a brilliant play of iridescence, usually in red, green, or
blue.) Alex Ritchie, curator of p a l e o n t o l o g y at the Australian M u s e u m in
Sydney, came up with the idea of raising the money on national television,
and $340,000 was donated by individuals and companies to purchase the
precious fossil and keep it intact.
"Eric" has now been assigned to the genus Leptocleidus ("slender clavicle")
and probably represents a new species. T h e opalized "Eric" was found with
the
with
an
skeleton,
once looked
something like this.
Leptocleidus
about
was
tofeet long.
fish vertebrae inside the gut, suggesting a small-prey diet, and gastroliths that
m a y have aided in digestion or even been used as ballast. T h e other Leptodeidus
species (which all have proper n a m e s ) are less d r a m a t i c in substance; all are
smallish plesiosaurs with a triangular skull that has a distinct crest. T h e
nominal species (Leptodeidus superstes) was found in England (Andrews 1922a),
and representative fossils have also been found in S o u t h Africa (I., capettsis). A
western Australian specimen is L. demai, named for John Clema, who sponsored the expedition that excavated the fossils. Leptodeidus demai was the largest
species, reaching about 10 feet in length; the other species were about porpoise
sized, and their size and structure suggest that they might have been inshore
or even freshwater inhabitants. "Leptocleidids," wrote Cruickshank et al. in
1999, "are relatively small plesiosaurs which probably fed on fish and cephalopods in the surf zone or estuaries, and seem to be related to the 5 m long Early
Jurassic genus Rhomaleosaurus." T h i s has been borne out by O'Keefe's 2002
cladistic analysis, which shows that Leptodeidus is indeed a member of the
Rhomaleosauridae.
Another opalized plesiosaur skeleton was found in 1967 by John and M o l l y
A d d y m a n , opal miners from A n d a m o o k a , South Australia. In November
2000, the "Addyman plesiosaur" was bought by the Adelaide newspaper the
Advertiser for $25,000 and donated to the South Australian M u s e u m . It was
studied by the museum's paleontologist Ben Kear (he is also examining an
ichthyosaur skull to see if the animal was d e a f ) , who assigned it to the genus
Leptodeidus, but he did not identify the species. T h e skeleton, which is about 80
percent complete, is just over 2 feet long. Because it was "by far the smallest
and most immature example of the genus yet discovered," Kear realized that it
was a baby, or, in paleo-speak: "Small size coupled with incomplete fusion of
the basicranial elements, vertebral centra, neural arches and cervical ribs, and
poorly ossified articular surfaces on humerus and femur indicate that the
specimen is at an early stage in its ontogeny." But even for its small size, this
specimen (cataloged as S A M Pi5980) had very large flippers. Kear wrote:
T h e most distinct growth related change evident in S A M P15980 is the
disproportionately large size of the propodial elements, which equal
around 17.5% of the estimated 700 mm total body length. T h i s contrasts
with more mature specimens of Lcptocleidus spp., in which the propodials are
considerably smaller, generally representing only around 9 % 1 1 % . T h e
relative proportions of the propodial elements also appears to vary with
ontogenetic stage, juveniles showing very nearly equal humeral and femoral
lengths unlike mote mature individuals in which the femur is around 6%
larger. T h e functional implications of this disparity in l i m b proportions
might reflect the presence of differing locomotion and buoyancy regimes
between adult and juvenile individuals. Indeed Wiffen et al. (1995) suggested that juvenile plesiosaurs m i g h t have specifically employed slower
swimming speeds and a more hydrostatic (passive) regulation of body
trim. In the case of taxa such as Lcptocleidus, in which both juveniles and
adults appear to have exclusively occupied freshwater to shallow near-shore
marine dcpositional environments, alternative behavioural strategies such
as migration, differing prey preference a n d / o r feeding zones within the
water column might also have been present to avoid intraspecific competition between age groups.
One of the biggest and most formidable of the pliosaurs was Kronosaurus,
from the early Cretaceous of Australia. ( T h e name comes from Kronos, the
Greek mythological figure that ate his children.) In 19311932, an expedition
from the M u s e u m of Comparative Z o o l o g y of Harvard University, headed by
W. E. Schevill, discovered an almost complete skeleton of Kronosaurus in the
A r m y Downs region of Queensland. It was dynamited out by Schcvill's
assistant, and about four tons of rock and fossil was wrapped in bloodied
sheepskins and sent back to Harvard. In the 1950s, renowned paleontologist
Alfred Sherwood R o m e r helped m o u n t the new Queensland material in
Harvard's M u s e u m of Comparative Zoology, but the bones were badly
eroded and required much plaster and even more imagination to reconstruct
the skeleton. ( C y n i c s referred to it as "Plasterosaurus" at the time, because so
much of the original fossil material was encased in plaster.) As assembled by
Romer and his colleagues, Kronosaurus had 43 presacral (forward of the pelvis)
vertebrae, which stretched it to a length of 42 feet, the size of a humpback
whale. Subsequently unearthed pliosaur fossils substantially more complete
than "Plasterosaurus" have had no more than 35 dorsal vertebrae, m a k i n g
Romer's original estimate somewhat suspect, and reducing the Harvard Kronosaurus to a more modest 35 feet. (Despite the revisionists, however, in 1999,
C r u i c k s h a n k et al. wrote, " K r o n o s a u r u s queenslandicus is a giant form reaching
nearly 14 m [45 feet] as shown by the reconstructed skeleton on display at the
Harvard M u s e u m of Comparative Zoology.") T h e massive, crested skull of
the Harvard specimen measures no.7 inches ( m o r e than 9 feet) long, which
makes it more than twice as long as that of Tyrannosaurus, the largest of the
terrestrial carnivores, whose skull has been measured at 52 inches. (Both
wither in comparison to the skull of today's s p e r m whale, which can be i&Jeet
l o n g . ) Even at the reduced length of 35 feet, the early Cretaceous Kronosaurus
was one of the largest marine reptiles and certainly one of the most terrifying
predators that ever lived. Its massive jaws held an array of teeth that were u
inches long, longer than the canines of the Pleistocene saber-toothed cats,
and equaled today only by the lower jaw teeth of sperm whales.*
T h e type specimen for Kronosaurus queenslandicus is based on a jaw fragment
with six teeth that was discovered by Andrew C r o m b i e in 1899 near H u g h e n den in central Queensland and described by Heber L o n g m a n in 1924, 1930,
and 1932. In 1929, more material was found at the same location, and under
Longman's supervision (he was the director of the Queensland M u s e u m ) , a
restoration of the specimen was attempted ( L o n g m a n 1932). T h e skull is
broad, low, and flat very different from the Harvard specimen, which is
deeper and more robust and has had an unwarranted sagittal crest applied to
it in plaster. It is likely that it is from a different species, although both are
still classified as K. queenslandicus. In 1998, Australian paleontologist John Long
wrote, " M o l n a r (1991) doubts that the Harvard skeleton is really the same
species as the type material described as Kronosaurus queenslandicus by Longman
(1924) since the two specimens came from different aged strata." As Tony
T h u l b o r n and Susan Turner (1993) wrote, "it is difficult to judge the extent of
* Of course, there are teeth longer than n inches today, such as the ivory tusks of walruses
and elephants and the spiraling tooth of the narwhal, but none of these are used for biting.
In fact, it is questionable whether the sperm whale bites anything with its massive ivory
pegs; most of the squid eaten by the teuthophagous sperm whales show no sign of having
been bitten, and it is now believed that the whales send out focused bursts of sound that
stun or even kill the squid, which the whales then gobble up.
The
30-joot-long
Kronosaurus
quecnslandicus,
any differences in skull shape, given that a considerable part of the Harvard
known from
early
thing that "we still do not have a description of the skeleton, despite its
relative completeness" ( L o n g 1998).
Colin M c H c n r y of Canowindra, N e w South Wales, has a thing about
the
Cretaceous
Australia,
oj the
most
fearsome
massive,
me in 2002, he said:
of
was one
long,
jlat-lopped
more than
also doesn't have that crest, or huge bulbous protrusion on the top of the
r e x , the largest oj
all
with big flippers, Kronosaurus was more of a sea lion with the skull of a croc.
carnivores.
terrestrial
Liopleurodon
was the terror oj the
late furassic seas of
Europe.
missing only the tail, and with nineteen dorsal vertebrae, it was estimated to
have reached a total length of 30 feet. At this length, Kronosaurus boyacensis
would have been a fearsome predator; its sharp, ribbed teeth were the s i z e ol
bananas.
T h e gigantic pliosaur Kronosaurus ( o f whatever species) has been implicated in a direct attack on the elasmosaur Woolungasaurus. W h e n Australian
paleontologists T h u l b o r n and Turner (1993) examined the skull of the elasmosaur found in Queensland in 1980, they saw that it was crushed so badly
that no one had noticed the tooth marks. (Indeed, the skull was in such bad
shape that it was actually broken in half, and the two pieces were sent to two
different institutions, the Queensland Geological Survey and the Australian
M u s e u m in S y d n e y . ) T h u l b o r n and Turner described some of the damage to
the reassembled skull as "not readily explicable as the results of post-mortem
crushing and distortion" and concluded that it had been inflicted by a predator with exceptionally large teeth, probably Kronosaurus. Of the possible encounter, they wrote:
T h e long and extremely flexible neck of the clasmosaurs, sometimes comprising more than 70 vertebrae, was probably an adaptation fot seizing
fast-moving fishes and cephalopods; the head could be swept smoothly
and rapidly through a wide range, both sideways and dorsovcntrally [up
and d o w n ] , in pursuit of such elusive prey. . . . at the same time, this long
(also called Walking with Dinosaurs), Dave Martill and Darren Naish wrote Walking with
Dinosaurs, The Evidence: How Did They Know That? In answer to the question "How big was
Liopleurodon and
the
ichthyosaur
Opbtbaltnosaurus:
He [Liopleurodon] raises his massive head slowly and then drives his flippers
down. As he lurches forward, ammonites are sent tumbling through the
water and fish arc dragged off the coral in his wake. H i s mouth opens and
snaps firmly shut round the m i d - p o r t i o n of the struggling O p h t h a l m o saurus. T h e power of the attack carries both his head and his prey clean
Liopleurodon," they wrote: "Because it is not possible to simply put whales onto weighing
scales, experts disagree over the weights of these animals. Some say that the largest blue
whales may reach an astonishing 2 0 0 tonnes, while others say that they probably don't even
reach 1 0 0 tonnes. Regardless, weights within the range were then applied to Liopleurodon.
However, most of a whale's bulk is carried in the thick blubber layers it carries for use on its
long migrations, and to insulate it from the cold of the polar seas it often frequents.
Liopleurodon was a denizen of warm tropical seas and would not have had such blubber. We
therefore estimate that even the biggest pliosaurs would not have weighed as much as the
biggest whales."
out of the water, where they hang for a moment before he brings both
Skeletal
down with explosive force. A m o n g all the spray and blood, his victim dies
reconstruction
instantly, her body punctured by his long teeth and her back broken. He
adjusts her limp corpse in his mouth, repeatedly biting and shaking it.
Eventually it breaks into three pieces and, grasping the front portion, he
rises to the surface, flicking to the back of his gaping pink throat and
Liopleurodon
(after
Newman
Tarlo
196y).
and
Notice
swallowing.
Although most of the skeleton was destroyed in the process of removing it,
a Russian Liopleurodon was discovered on the right bank of the Volga River in
1938. T h e remainder, which consisted of the skull and pectrum, was saved and
described in 1948 by the Soviet paleontologist Novozhilov as Pliosaurus rossicus.
Then the ubiquitous Beverly Halstead (1971) recognized it as Liopleurodon and
renamed it Liopleurodon rossicus. In 1966, Ken Carpenter described a somewhat
smaller version of Liopleurodon, which he named Plesiopleurodon ("near Liopleurodon"), found in the Belle Fourche Shale of W y o m i n g . Like its larger namesake,
Plesiopleurodon had powerful jaws and eight pairs of caniniform teeth, which
were smooth and not striated like those of other, similar species. (In his
thesis, N o e [2001] says that Plesiopleurodon is different enough from L.Jerox to
suggest that it might belong to another genus altogether.)
In 1984, a fragmentary skeleton was discovered by a student near the village of Aramberri, in Nuevo Leon, northwestern M e x i c o . It was originally
thought to have come from some sort of dinosaur, but later examination
revealed it to have belonged to a pliosaur. Eberhard ( " D i n o " ) Frey, Celine
Buchy, and Wolfgang Stinncsbeck examined the material in the m u s e u m in
Linares, Mexico, and discussed it in a presentation at the European W o r k -
ribs
(gastralia).
Braekiosaurus
at
78
animals could not possibly support themselves on land and had to spend most of their lives
in the water, but this idea has now been thoroughly rejected.) The weight of a full-grown fin
whale is about 76 tons, and right whales and bowheads regularly go over 1 0 0 tons. The blue
whale, which Lyall Watson ( 1 9 8 1 ) calls "the largest animal the world has ever known," weighs
in at well over 150 tons; Watson reports the "recorded maximum of 1 7 8 . 0 0 0 kg ( 1 9 6 tons)."
Brachauchenius
("short neck"in
Creek) was the
shortest-necked
plesiosaur known,
with only eleven
cervical vertebrae.
Based on a j-footlong skull,
Brachauchenius
is estimated to have
reached a length of
j6'Afeet, making 11
lining reduced drag, hydrofoil l i m b s p e r m i t t e d motion through a dense, yet
great mega-predators
of the late
a powerful bite, sharp teeth and powerful neck made pliosaurs top predators
in M e s o z o i c food webs." T h e narrow, elongated skull and widely spaced
sharp teeth suggest that Peloneustes was a fish eater; with its wide skull and
powerful (often broken) teeth, Liopleurodon obviously favored large, hardboned prey.
Another of the more formidable giant pliosaurs was Brachauchenius lucasi,
identified by Samuel W i l l i s t o n in 1903 from a skull and vertebrae found in the
Benton Formation of Ottawa County, Kansas. He called it Brachauchenius
("short neck" in Greek) because it was "the shortest-necked plesiosaur
known" at the time, with only eleven cervical vertebrae in a neck that was
about 75 percent as long as the skull; lucasi was for Frederick A. Lucas of the
U.S. National M u s e u m , "who has done much valuable work in American
Cretaceous.
paleontology." T h e short neck and relatively long skull are pliosaurian features, and the skull is distinctive for its broad, triangular, mosasaur-like shape
that ends in a point, unlike that of other large pliosaurs such as Liopleurodon
and Pliosaurus, whose skulls taper into a narrow, blunt snout. T h e large teeth
have striations that branch toward the root, unlike the straight grooves on the
teeth of Jurassic pliosaurs. Known from three nearly complete skulls with
mandibles and two partial skeletons, and based on a 5-foot-long skull,
Bracbauchenius is estimated to reach up to 56 /2 feet in length and is therefore
!
one of the last of the great mega-predators, dating from the CenomanianTuronian (early late Cretaceous). T h e s e gigantic forms are younger than the
kronosaurs and pliosaurs and may have evolved possibly from unknown
short-necked clasmosaurids to fill the ecological niche left vacant by the
disappearance of the earlier gigantic species of the Cenomanian and early
Turonian. An alternative explanation is that Brachaucbenius is actually a shortnecked pliosaurid. After the first Bracbauchenius lucasi from Kansas, a second
specimen was collected from the Eagle Ford Formation near Austin, Texas,
and was described by W i l l i s t o n in 1907. A third specimen, more complete and
somewhat better preserved, was collected from the Greenhorn Limestone in
Russell County, Kansas, and is on display at the Sternberg M u s e u m in Hays,
Kansas.
Judy M a s s a r e has studied the eating habits of the plesiosaurs, but she does
not attempt to resolve the bothersome question of how these gigantic reptiles
could maintain enough energy in the cold ocean to s w i m so fast and attack so
viciously. In her 1987 analysis of the "tooth m o r p h o l o g y and prey preference
of M e s o z o i c marine reptiles," she wrote that "the teeth of many plesiosauroids, such as Plesiosaurus dolichodeirus, P. brachyplerygius, Muraenosaurus leedsi, and
Cryptoclidus eurymerus, are very long, slender cones with sharply pointed apices";
they were similar to the teeth of ichthyosaurs, in that they rarely show wear
and were probably used to pierce soft prey. S h e compares the sharp, ridged
teeth of Liopleurodon to the teeth of killer whales, saying that they are frequently broken and worn, "suggesting a diet of fleshy prey with fairly large
bones, such as very large fish and other reptiles."
In a discussion published in 1959 ("Pliosaurus bracbyspondylus [ O w e n ] from
the Kimmeridgc C l a y " ) , Tarlo wrote:
In the centre aisle of the University M u s e u m , Oxford, there is on exhibition a giant mandible belonging to a Pliosaur from the Kimmeridge C l a y
of Cumnor, Berkshire. It was first noted by Prestwich and seems to have
been acquired by the M u s e u m some time between 1880 and 1888. In 1933
M r . H. J. H a m b i d g e completed the long and arduous task of renovating
this specimen which he had first known in 1907. Professor W. J. S o l l a s had
intended to describe the m a n d i b l e in 1936, but unfortunately he died the
same year. Since that time, this remarkable mandible has remained unidentified and undescribed and no recognition has been given to the skilful
work of M r . H a m b i d g e .
Tarlo then describes the mandible, which, if complete, would have been 9%
feet long. " W i t h o u t doubt," he wrote, "it belongs to the largest pliosaur ever
recorded, somewhat exceeding the size of the Cretaceous Kronosaurus." (In
1959, Kronosaurus was believed to have been 45 feet l o n g . ) Tarlo wrote that the
giant pliosaurs had previously been l u m p e d into Pliosaurus macrotnerus, but "a
detailed examination of these remains has demonstrated the existence of two
different Pliosaur genera represented by the species Pliosaurus bracbydeirus and
Stretosaurus macrotnerus. (Stretosaurus was the genus that Tarlo erected for the
newly described material, but the name would not last l o n g . )
W h e n Tarlo examined pliosaur fossils from the Kimmeridge C l a y formation in Ely ( W i l t s h i r e ) , he found that the Kimmeridgian pliosaurs could be
separated into two groups: one contained Pliosaurus brachyspondylus and P.
bracbydeirus, and the other was a new genus that he proposed to call Stretosaurus,
after the village of Stretham, where the giant pliosaur fossil had been d i s covered in 1952. Further study, however, revealed that the bone he took to be a
scapula was actually an ilium, which meant that his analysis was wrong, so he
made Stretosaurus a junior s y n o n y m of Liopleurodon. In his 1989 study, Halstead
wrote, " T h e basis for the erection of a new genus likewise foundered and the
name Stretosaurus is now redundant and should be replaced by the longestablished Liopleurodon." (In his 1959 paper on Stretosaurus, Tarlo suggested that
this giant pliosaur might have been even larger than Liopleurodon, but now that
the two genera have been synonymized, Liopleurodon retains the title of largest
pliosaur k n o w n . ) Again employing the peculiar approach that resulted from
"mcgaplcurodon," but they insist that this is just a n i c k n a m e and should have
no scientific standing.
Another large pliosaur found in the Oxford C l a y is Simolestes ("snub-nosed
robber") vorax ("voracious"), which reached a length of 20 feet maybe more.
T h e ends of its jaws were expanded into a spatulate rosette a r m e d with huge
caniniform teeth, which suggested a powerful bite-and-twist feeding style:
" T h e rosette of symphysial teeth was probably used for tearing large chunks
of llesh from its prev, or for biting chunks out ot larger a m m o n i t e s "
Martill
late, tooth-studded rosette is likely an artifact of crushing and did not exist in
life in any pliosaurs.) Maresaurus is known only from a yard-long skull and a
few cervical vertebrae, but its 20-foot-long body was probably like that of
other pliosaurids. (According to Noe, Simolestes and Maresaurus might be synonymous, and both might be synonyms for an older taxon called Eurysaurus.)
T h e r e were gigantic pliosaurs in Jurassic N o r t h America too. In 1898,
W i l b u r Knight found a fossil he christened Megalneusaurus, or "great swimming
lizard," and described it as "the largest known animals of the Sauropterygia."
Its forelimbs were said to be 7 feet long ( W i l l i s t o n 1903), but the actual
remains appear to have been lost. Recently, a smaller specimen was found in
southern Alaska ( W e e m s and Blodgett 1996), but the classification of this
specimen remains unresolved. According to Creisler, Robert Bakker is currently studying the material to provide a more detailed, updated description.
Although the skull was not preserved, Bakker has suggested that the animal's
head may have been n feet long at least based on the Harvard reconstruction of Kronosaurus. T h e fossils of Megalneusaurus were found in the upper part
of the late Jurassic S u n d a n c e Formation of W y o m i n g . W i l l i s t o n (1903) wrote,
"A large portion of the type species is known; the parts so far described are the
vertebrae and limbs." However, some of the original remains (ribs, vertebrae)
mentioned by Knight and W i l l i s t o n have apparently been lost, since the
surviving specimen consists only of a forepaddle, some vertebrae, and fragments of a pectoral girdle material that many researchers (but not a l l )
consider inadequate to diagnose a genus and species. "A full-grown Megalneusaurus m a y have been in the 3540-foot range or larger," wrote Creisler, "but
such estimates should be greeted with caution until more hard facts are
known."
W h e n workers on U.S. H i g h w a y 81, south of Concordia, Kansas, came
across some fossilized bones in a road cut, they carefully dug the material out
and turned it over to the University of Kansas M u s e u m of Vertebrate Paleontology. T h e fossils were described in a 1944 publication by Elmer S. R i g g s .
He had "the skull with mandible, fifty vertebrae, many ribs, most of the
pectoral girdle . . . and the ischia almost entire" enough to recognize it as a
new species of plesiosaur that he named Trinacromerum ("three-tipped femur")
willistoni (after Samuel W i l l i s t o n ) . T h e genus Trinacromerum had been named in
i888 by Cragin and was applied first to T. bentonianum, named for the Benton
Formation in Kansas where it was found. Riggs's paper consisted entirely of a
description of the fossil (e.g., " T h e sutures joining the pedicles and the
cervical ribs to the centra are plainly m a r k e d " ) , and there was no speculation
as to what Trinacromerum looked like. In 1996, in response to a tendency to
make Dolichorhynchops ("long snout face") a synonym for Trinacromerum, Ken
Carpenter wrote that "the p o l y c o t y l i d s Trinacromerum and Dolichorhynchops are
separated by autapomorphies in the skull of Dolichorhynchops," m a k i n g them
separate genera after all. As shown by a life-size model on display in the
Sternberg M u s e u m , Trinacromerum was a small, short-necked plesiosaur (a
pliosaur), approximately the size of a m o d e r n sea lion. It is shown on the
beach, suggesting ( s u r p r i s i n g l y ) that it might have been able to come ashore.
S. P. Welles (1962) wrote, " T h e nomcnclatorial confusion surrounding the
short-necked U p p e r Cretaceous plesiosaurs from N o r t h America has been
resolved by eliminating the names Piratosaurus and Polycotylus as nomina vana.
Our concept is based on the first adequately known genus, Dolichorhynchops."
T h e Polycotylidac were the last of the short-necked pliosaurs. In some classifications they are included under the Pliosauridae. T h e y were generally
smaller than the pliosaurids proper, averaging about 10 feet long. T h e head
was large, at least as long as the neck, and the snout was generally very
elongate. T h e y were short-necked, with uniform conical teeth and none of
the massive caniniform teeth that characterized the other pliosaurs. According to Dawn A d a m s (1997), Trinacromerum bonneri was the "last and fastest
pliosaur of the Western Interior Seaway." Its flippers were "the longest wingfins known," equal in length to its dorsal spinal column, enabling the animal
to achieve unprecedented speed in the water. ("Pliosaurs," she wrote, "have
always been regarded as particularly high-speed swimmers . . . highly mancuverable animals, capable of changing direction skillfully in pursuit of large
prey.") A "tongue and groove" articulation of the digits "further increased
wingfin strength and rigidity along the longitudinal axis and m i n i m i z e d
torsion of the wingfin as a whole, which permitted the development of longer
wingfins with more wingloading and greater propulsive power."* Bakker
For this paper, Adams also d a w a reconstruction of Trinacromerum bonneri that might have
What does
Trinacromerum
remind you of? A
10-foot-long, fourflippered penguin? It
(1993b) wrote that this type of long-snouted pliosaur (called dolichorhynchopine because of Dolicborbyncbops) became the most common short-necked
plesiosaurs in the Western Interior Seaway after the extinction of the ich-
thyosaurs at the end of the Jurassic period. Indeed, they began to resemble
ichthyosaurs, with their enlarged eye sockets, longer jaws, and reduced teeth.
pliosaurs, becoming
extinct in the late
been better conceived. The proportions arc commensurate with her description, but for
Cretaceous, about
some reason, she chose to picture the animal as if it were chrome-plated, which, if correct,
70 million years
would probably have cut down water resistance and increased its not inconsiderable speed
ago.
even more, but it raises a whole new set of questions about the integument of pliosaurs.
T h i s suggests that they filled the gap left by the departing ichthyosaurs, which
were also fast-swimming ambush predators.
W h e n Cruickshank, M a r t i l l , and N o e (1996) examined the ribs of a
previously undescribed pliosaur fossil found in 1994 in the Oxford Clay
( m i d d l e Jurassic) of Cambridgeshire, England, they named it Pachycostasaurus
because the bones were much heavier than those of other pliosaurs a characteristic known technically as pacbyostosis. T h i s condition is known today in
manatees, animals that spend much of their time grazing on the bottom, so it
was suggested that Pachycostasaurus dawni (for Alan Dawn, who found the
fossil) was, like the manatees, a bottom-feeder. No gastroliths were found in
conjunction with the 10-foot-long fossil, suggesting that its dense bone structure may have been sufficient ballast to allow it to sink and feed on or near the
bottom. T h e teeth of Pachycostasaurus were striated, sharp, and conical, which
indicated predacious feeding, but because the skull was comparatively light
and delicate, the authors suggested that this large pliosaur probably d i d not
attack big, strong prey animals. T h e y wrote, " W e speculate that Pachycostasaurus fed on nektobenthic arthropods, cephalopods, or possibly on the
assumed nektobenthic, heavily scaled g a n o i d fishes. Pachycostasaurus might have
fed on mid-dwelling, soft-bodied prey that didn't put up much of a struggle,
such as burrowing shrimps."
Pachycostasaurus was one of the p r i m a r y subjects of N o e s 2001 study (the
others were Liopleurodon and Sitnolestes), and in his thesis, he modified his earlier
description of the hunting strategies of this dense-boned creature. He wrote,
"Pachycostasaurus is here interpreted as preying on hard-boned vertebrate prey,
the exceptionally stout and heavily ornamented teeth indicating prey even
more difficult to subdue than that tackled by Liopleurodon." T h e only Pachycostasaurus fossil was 10 feet long, much smaller than Liopleurodon (which may have
been 50 feet l o n g ) , and it may have been a juvenile. But thus far, it is the only
specimen known, and, wrote Noe, the "definitive interpretation of preferred
prey will have to await the discovery of further examples of Pachycostasaurus."
M c H e n r y (personal c o m m u n i c a t i o n 2002) has attempted to resolve the
confused and confusing state of pliosaur systematics by recognizing five
major groups, or "families." T h e y are the Rhomaleosauridae, Pliosauridae,
Simolestes,
Maresaurus,
PMegalneusaurus,
Plesiopleurodon,
PPolyptychodon.
Middle
From the late Jurassic to the early late Cretaceous, giant pliosaurs were the
terrors of the seas. Like all members of the Plesiosauria, they had heavily constructed bodies, short tails, and four powerful paddles used to s w i m and steer.
pliosaur
Pachycostasaurus dawni
with a short neck and a large head. T h e real evolutionary story may be more
complex, and some researchers now think that the pliosaur design may have
on
soft-bodied prey
Pliosaurus,
Liopleurodon,
Bracbauchenius,
Polyptycbodon,
Kronosaurus had
huge
skull with the biting power of killer whales and crocodiles. Exactly how large
these great predatory sea dragons grew remains an intriguing question. T h e r e
is no question, however, that these prehistoric reptiles are the quintessential
sea monsters; twice as long and ten times as heavy as the largest living
crocodiles, the pliosaurs were probably the most terrifying marine predators
that ever lived. T h e y dominated the seas the way the carnivorous dinosaurs
dominated the land. Were it not for the fossil evidence that unquestionably
demonstrates their existence, they would surely be relegated to the realm of
nightmares.
as
mud-dwelling,
such
burrowing
shrimps.
Pachycostasaurus
was about
to feet long.
The Mosasaurs
Although there are mosasaur fossils aplenty in what was once a vast inland sea
in N o r t h America, the first mosasaur fossil was found in 1780 in Maastricht,
the Netherlands. Workers in a limestone mine 90 feet deep discovered a huge
fossilized skeleton of a sort that had never been seen before. An army surgeon
named C. K. Hoffmann directed the q u a r r y m c n to bring the whole rock containing the fossil to the surface, but while he was trying to extricate the fossil
from the matrix, a clergyman named Goddin, who owned the land in which the
mine was sunk, sued him and won possession of the rock-bound monster. He
also got Dr. Hoffmann's money, because the unfortunate surgeon was m a d e
to pay the costs of the legal action as well. Goddin built a chapel to house the
fossil, but during the 1795 siege of M a a s t r i c h t by Napoleon's army, it mysteriously disappeared. W h e n it was located (it had been "liberated" by N a p o leon's grenadiers), it was sent to Paris, where various people argued about
what it was. Because of its size, Pieter Camper, a renowned Dutch anatomist,
Found in l~]8o in
Maastricht, the
Netherlands,
Mosasaurus
liollnunni was
the first mosasaur
everfoundand the
largest, reaching 58
feet. The enormous
size of this animal,
along with its
powerful jaws and
teeth, meant that
almost any creature
believed it was a toothed whale; French scholar Barthclmy Faujus dc SaintFond ( k n o w n as Faujus) published an elaborate description in which he
compared it with a crocodile. Unimpressed with its size, Pieter Camper's son
Adriaan Gillcs C a m p e r correctly compared it with a varanid lizard, and based
including hard-
mosasaurs.
and 1791 and followed in several later publications," but because he was so
much better known, Cuvier is usually given credit for correctly identifying the
mosasaur as a lizard. In 1820, S a m u e l S o m m c r i n g suggested the name Lacerta
gigantea for the M a a s t r i c h t mosasaur, but it turns out that he was describing
History Museum of Maastricht and members of the Dutch Geological Society revealed
that the skeleton, dispersed over an area of more than 40 square meters, included portions
of both lower jaws and parts of the upper jaw, as well as vertebrae and ribs. With the
exception of the missing tail bones, the skeleton is more or less complete,
t In
Cope's rule include the thcropod dinosaurs (which culminated in T. rex) and also camels,
elephants, and horses. Bonner points out that N. D. Newell ( 1 9 4 9 ) also showed that
"the same principles apply to invertebrates [including] foraminiferans, arthropods, echinodcrms. brachiopods and ammonites, all widely separated groups." But then Bonner says
that "a close, more finely tuned inspection of the fossil record shows that there is as much
getting smaller as there is getting larger," which would indicate that Cope's rule is tautologically applicable only in those taxa where there is a demonstrable increase in size in the
fossil record.
for Plioplatecarpus bouzeaui, which had the best binocular vision of any mosasaur.
T h e enormous size of this animal, along with its powerful jaws and teeth,
meant that almost any creature was potential prey, including hard-shelled
turtles and probably even other mosasaurs.
Around 1829, a mosasaur fossil was found by Major Benjamin O T a l l o n
near the Great Bend of the Missouri River, between Fort Lookout and Fort
Pierre in what is now central South Dakota. He transported it to his garden in
St. Louis, where it was seen by Prince M a x i m i l i a n of W i e d , who bought it
from O'Fallon, brought it back to Germany, and placed it in the museum at
Bonn. In 1845, German paleontologist August Goldfuss described it and
named it Mosasaurus maximiliani, after his patron. In 1830, when Richard Harlan
was given the end of the snout, he believed that it came from an ichthyosaur,
so he named it Ichthyosaurus missouriensis. In the section on mosasaurs in Water
Reptiles of the Past and Present, W i l l i s t o n (1914) mentioned Harlan's misidentification, and wrote that "some time previously, it has since been found, some
fragments of the same species were described by Harlan, an American author,
under the name Ichthyosaurus missouriensis." But in his 1967 monograph on
American mosasaurs, Dale Russell lists M. missouriensis as one of the mosasaur
species from the Pierre Shale and identifies three other specimens, one from
M o n t a n a and two from S o u t h Dakota. T h e end of the snout that Harlan had
mistaken for part of an ichthyosaur was likely a missing piece of the O'FallonGoldfuss mosasaur, so M. missouriensis can take its rightful place as a proper
mosasaur after all.*
In 1899, Louis Dollo described another species of mosasaur that had been
found in the vicinity of M o n s in Belgium, some distance from Maastricht but
from the same U p p e r Cretaceous formation as the original Mosasaurus. According to L i n g h a m - S o l i a r and N o l f ' s 1989 description of Prognathodon (originally named Dollosaurus), it was a 14-foot-long mosasaur similar in shape to
another mosasaur called Plioplatecarpus. T h e forward portion of the upper jaw
( p r c m a x i l l a ) was a r m e d with a clublike arrangement of four teeth, which
looked like a four-fingered paw and was responsible for the name Prognathodon
" In fairness to Harlan, he later recognized that the piece had come from a mosasaur, and
after showing it to Richard Owen, he tried to change its name to Ratrachiosaurus ("frog-like
lizard"), but the name was officially changed to Mosasaurus missouriensis in an 1 8 3 9 publication.
but have to come ashore to lay their e g g s . ) T h e y did not enter a realm devoid
of predators, for the ichthyosaurs and plesiosaurs had colonized the oceans
long before the mosasaurs did, but the ichthyosaurs were in decline by the time
the mosasaurs arrived. T h e disappearance of the ichthyosaurs may be one of
those mysteries of extinction with no answer, and it may have been nothing
more than an evolutionary coincidence. Dale Russell (1967), however, believes
there is a connection: " T h e remarkable convergence of some of the later mosasaurs with primitive Triassic ichthyosaurs shows that they were an ecological
replacement of the declining ichthyosaurs." " S o m e mosasaurs may have been a
belated ecological replacement of Triassic ichthyosaurs," wrote L i n g h a m Soliar in a letter to me, "but not of the Late Cretaceous thunniform ichthyosaurs. Triassic ichthyosaurs were ambush predators, and gave rise to pursuit
predators at the beginning of the Jurassic, all long before mosasaurs. T h e
vacated Triassic ichthyosaur ecological niche was partially filled by marine
crocodiles and much later by aigialosaurs. T h e n mosasaurs came along and
did the job of ambush predation better than the others put together and some
may have filled a similar ecological niche to that of Triassic ichthyosaurs."
W h a t e v e r they were, the ancestors of mosasaurs have not been identified,
but they probably looked a lot like aigialosaurs. In their 1992 survey of these
mid-Cretaceous varanid lizards, Robert Carroll and Michael DeBraga noted,
"we know almost nothing of the history of lizards between the end of the
Jurassic and the Late Cretaceous, by which time most of the modern families
had differentiated." T h e exception is the aigialosaurs, terrestrial lizards 3 feet
long or more, with a tail as long as the head and body combined, not unlike
t o d a y s monitor lizards ( V a r a n i d a e ) . T h e r e are several nearly complete skeletons, representing five species: Aigialosaurus dalmaticus, A. novaki, Opetiosaurus
buccicbi,
Carsosaurus marchesetti,
and
Proaigialosaurus hueni,
all
similar enough
to
suggest that they might belong to the same species. However they are classified, the aigialosaurs were probably part of the ancestry of the mosasaurs. But,
as L i n g h a m - S o l i a r (1994c) wrote, "Aigialosaurs demonstrate many conditions
that might make them a suitable ancestor for the Mosasauridae, but there are
problems: for instance, aigialosaur material is scanty and aigialosaurs and
mosasaurs are both known only from U p p e r Cretaceous deposits. T h i s poses
problems regarding an aigialosaur ancestry unless it could be established that
Aigialosaurs
smallish
were
terrestrial
today's
monitor
lizards.
they had arisen earlier." As Carroll and DeBraga (1992) wrote, " N o significant
ancestors
characters of the skull are known that distinguish aigialosaurs from primitive
mosasaurs
The
of
(and
varanid
advanced varanids and aigialosaurs from the more primitive genera now
included in the Varanidae."
lizards)
From the very limited fossil record (all aigialosaur material comes from
Yugoslavia except Proaigiahsaurus, which was found in the Solnhofen limestone
of Bavaria), it appears that even if aigialosaurs cannot be identified as ancestral mosasauroids, then something very similar began the mosasaur line,
which Gordcn Bell (1997a) called a "27 millionyear procession of vertebrate
evolution so complete that it may well rival the example provided by the fossil
record of horses." T h i s m a y be exaggeration for emphasis. Although we do
have a Fairly good chronology of mosasaur development, the comparison of
mosasaur ancestry to that of horses might not be entirely justified. T h e fossil
record for horses shows replacements over time along natural lines of descent,
where one can see an increase in size, increase of speed through modification
of the limbs, elongation of the head and neck, and so on, but no horse species
has actually been shown to be ancestral to any other.* T h e r e are still many
unresolved problems with mosasaur ancestry and mosasaur phylogeny. It is
more than likely that the mosasaurs were highly marine aigialosaur-like animals. One might even say that aigialosaurs were m o n i t o r lizards caught in the
act of becoming mosasaurs. ( A l t h o u g h the mosasaur skull was like that of the
monitor lizards, with a joint in the m i d d l e of the lower jaw, the mosasaurs had
all died out by the end of the Cretaceous, so the marine mosasaurs are not
ancestral to the monitors.)
T h e fossil record is, as paleontologists term it, "scrappy," meaning that
their research is restricted to the occasional fossil that appears in a serendipitously uncovered layer of shale or sandstone. T h e r e is deeply buried
evidence of mosasaurids that no human eyes will ever see; therefore, paleontologists have to postulate a family tree with a lot of the branches missing.
Still, enough fossilized mosasaur pieces have turned up to enable scientists to
* Colin McHenry (personal communication 2 0 0 2 ) wrote, "In fact, it is almost paleontological dogma that we will never know what the ancestor of any animal looked like. For all
the fossils we have found, we have probably never found a fossil that was ancestral to
anything else. The odds against it are just too long for anything represented by a fossil to be
an actual ancestor. But fossils do represent relatives of the ancestor, and show us what the
ancestors probably looked like thus they make good models for the elusive ancestor. So we
arc able to reconstruct, sometimes quite confidently, what the ancestor of a certain group
must have looked like."
then, with the K - T extinction event, they abruptly vanished. By then, the
ichthyosaurs were long gone, and the pliosaurs were already in decline, so the
passing of the mosasaurs marks the end of an era as incredible as the reign of
the terrestrial dinosaurs: the time when reptiles ruled the seas.
By the mid-Cretaceous, some 90 million years ago, the continents were
arranged more or less where they are now, except that Australia was still
attached to Antarctica, and India, still attached to M a d a g a s c a r , had not yet
begun its northward journey. W o r l d w i d e sea levels had risen dramatically, and
much of N o r t h America and Europe was underwater, covered by the shallow
Tcthys Sea. At this time, the mosasaurs began the colonization of the shallow
waters that had been vacated by the ichthyosaurs. W i t h i n a geological m o ment, they had radiated to become the top predators of the Cretaceous seas.
T h e i r legs, originally equipped with feet designed for walking, developed
paddle-shaped flippers, and their tails lengthened and became flattened like
those of eels or crocodiles. In her 1994 analysis of the s w i m m i n g capabilities
of M e s o z o i c marine reptiles, Judy M a s s a r e wrote, "Not surprisingly, mosasaurs displayed m a n y adaptations for rapid acceleration that are characteristic
of ambush predators. T h e elongate shape resulted in a high surface-tovolume ratio for pushing against the water, and a relatively small frontal area
for pushing against it. . . . [ T h e ] expanded caudal area may have been an
adaptation for increasing thrust production in the distal part of the tail." T h e
tylosaurs had evolved a blunt prow on the end of the rostrum, which they may
have used as a battering ram to injure or incapacitate their prey. During the 25
million years that they prospered, the mosasaurs spread throughout the major
oceans of the world, ranging almost from pole to pole. ( T h e ichthyosaurs
lived for 150 m i l l i o n years, and the plesiosaurs for 140 m i l l i o n . )
M o s a s a u r s are characterized by a lizard-like skull in which there are two
openings behind the orbit. T h i s condition, known as diapsid, occurs in lizards,
snakes, and dinosaurs. T h e smallest mosasaurs, at a m a x i m u m length of 12
feet, are those of the genus Ciidastes ("locker," from the interlocked vertebrae);
they were the most lizard-like in form, with a thin, elongated body and a low,
triangular fin on the dorsal surface of the tail. Of all the mosasaurs, Ciidastes
had the shortest tail relative to body length, but also the best-developed tail
fin for marine locomotion, m a k i n g them more advanced than the ancestral
shallow water
inhabitant,
semiaquatic forms. T h e skull was short, the teeth smooth and sharp, and,
as Russell (1967) wrote, "the long, slender jaws of Clidastes were probably
adapted to rapid biting . . . and m i g h t have been effective in sawing a large
object into pieces of swallowable size." As for where in the water column
Clidastes hung out, there arc differences of opinion. W i l l i s t o n (1914) believed
that this species was a surface swimmer, and in their 1989 examination of bone
necrosis in diving mosasaurs, M a r t i n and Rothschild o p i n e d that Clidastes
probably lived near shore a n d " d i d not regularly engage in deep diving."
However, in a 1997 study of mosasaur bone microstructure, A m y S h e l d o n
found that Clidastes had particularly low bone density (a condition she calls
osteoporosis," which would have provided neutral buoyancy in deeper waters,
* In general usage,
osteoporosis
I'aehyoslosis,
condition of bones that arc denser or thicker than normal, a factor "largely confined to the
Sirenia and some extinct reptiles" (Domning and Buftrenil 1 9 9 1 ) . T h e term is also used f o r
pachycephalosaurs, ornithischian dinosauts that had a thick bony dome on top of the skull,
probabh used in intraspecihe head-bulling, . i s 111 bighorn sheep. I n personal c o m m u n i c a tions, various names have been suggested to describe a normal condition in which the bone
is unusually light, including
(from
tenuis,
pneumatic
elaphrosty
eiaphro,
tenuiosis
meaning "light in
mosasaurs,
reaching
rarely
Inject.
Skeletal recon-
and like Tylosaurus, it was likely a deep diver. ( N o matter how deeply a
struction oj the
mosasaur could dive, however, it had to come up for air; like all other reptiles,
giant mosasaur
Tylosaurus
proriger (after
Oshorn t8gg).
" T h e seas that rolled over Kansas in Cretaceous times," wrote Charles
Gilmore in 1921, "contained thousands of these a n i m a l s [mosasaurs] and in
the chalk bluffs of that region their remains are in such a state of preservation
that we are not only acquainted with their skeletal structure but with their
external appearance as well." A 25-foot-long Tylosaurus specimen found by
Charles Sternberg in Kansas in 1917 showed that "in life they were covered
with small, overlapping scales." In examining the specimen, Samuel W i l l i s t o n
detected "color m a r k i n g s " that consisted of "narrow, diagonally-placed parallel bars" (there was, however, no indication of the actual color). Like all
mosasaurs, Tylosaurus had an articular joint in the m i d d l e of each of the lower
jaws, which, combined with the very loose attachment at the front, allowed
these a n i m a l s to swallow large objects.
W o r l d sea level was at its highest during the M e s o z o i c , so there were vast
areas for the lizards to inhabit, and the remains that sank to the bottom were
preserved as fossils. T h e largest number of mosasaur fossils have been collected in Kansas, from the formation known as the N i o b r a r a Chalk. About
600 feet thick, the N i o b r a r a C h a l k extends from southwestern Kansas to
south-central M a n i t o b a , but it is best exposed in northwestern Kansas, where
b a d l a n d s have been cut along the bluffs of the S m o k y H i l l River and its
tributaries. It is composed of the compacted plates ( c o c c o l i t h s ) that are
remnants of the abundant, microscopic, golden brown algae ( C h r y s o p h y c e a e )
that lived in the w a r m , shallow sea. T h e upper portion of these deposits was
laid down between 87 and 82 million years ago during a period when the
Cope was probably a rylosaur, but its identification was based on fossil
material that could not be identified specifically, except that it belonged to some kind of
mosasaur. Russell
(1967)
included
T. dyspelor
Tylosaurus,
Tylosaurtis dyspelor
"Tylosaurus dyspelor
Tylosaurus proriger
vanum."
He seemed to think
could not find any features that clinched the match with certainty.
Nomen vanum
is an old
term meaning "empty name," no longer recognized by the International Code of Zoological
Nomenclature. T h e term now used is
nomen dubium
name thai has appeared 1 1 1 the literature nh a description but. according to other authors,
cannot be applied with certainty to a recognized taxon at a species level because the original
material lacks diagnostic features or, in some cases, is lost and cannot be reexamined
1 Creisler, personal communication).
their glistening teeth, and then the vertebrae and ribs. O u r delight was at
its height when the bones of the pelvis and part of the hind limb were laid
bare, for they had never been seen before in this species, and scarcely in the
order.
Tylosaurus proriger was originally named Macrosaurus proriger by Cope in 1869,
but Cope's arch-nemesis, O. C. M a r s h , renamed it Rhinosaurus and then Tylosaurus in 1872.* Proriger means "prow-bearing," referring to "the cylindrical
prolongation of the premaxillary bone beyond the teeth and a similar flat
prolongation of the extremity of the dentary" ( C o p e 1869c). One of the
largest of the mosasaurs, Tylosaurus proriger lived in the Niobrara Sea during the
late Cretaceous, some 85 m i l l i o n years ago. It was 20 to 50 feet long and had a
long, slim body; huge jaws; heavy, sharp, conelike teeth; and paddle-like fore
and hind limbs. It preyed on fish, shellfish, and probably smaller mosasaurs,
plesiosaurs, and hesperornithiform birds. Tylosaurus means "knob lizard,"
from the Greek tylos for " k n o b " and sauros for "lizard." T h e elongated cylindrical m u z z l e ( r o s t r u m ) projected beyond the front-most teeth in the upper jaw;
the tip of the snout was probably similar in function to the "ram" or "beak"
that the ancient Greeks and R o m a n s mounted on the prows of warships to
ram and sink enemy vessels. Tylosaurus might have used its "ram" snout to
stun prey, defend against sharks or other predators, or battle rivals of its
own species.
* This constant nomcnclatural revision is a handicap to those trying to write about the
fossils themselves. In order to chronicle the finds, one must identify the species that Cope
and Marsh named, which were often renamed by later workers. In such cases, a synonymy is
included in the later description, which enables the researcher to track the often convoluted
taxonomic history of a particular species. Here, for example, is Russell's 1 9 6 7 synonymy for
the mosasaur we now know as Tylosaurus proriger (the year following the paleontologist's
name represents the date of publication of the scientific name):
Macrosaurus proriger Cope 1 8 6 9
Macrosaurus pririger Cope 1 8 6 9
I.iodon proriger Cope
18691870
mosasaurs,
Tylosaurus
proriger lived in
the Niobrara Sea
during the
late
some
65
It
Cretaceous,
preyed
on fish,
shellfish,
probably
nepaeolicus, (Everhart writes that the name probably "comes from 'Ncpaholla,'
mosasaurs,
an earlier Indian name for the S o l o m o n River . .. meaning 'water on a hill.' ")
ichthyosaurs,
plcsiosaurs
brother of Charles), were described by C o p e (1874) and named Liodon nepaeolicus. Cope concluded that T. nepaeolicus was about a third smaller than the more
common T. proriger but was a separate species and not a juvenile. Everhart
concluded:
Since its description in 1874, Tylosaurus nepaeolicus has been considered to
be significantly smaller than T. proriger. T h e lack of complete specimens of
T. nepaeolicus seems to have discouraged further studies of this taxon. A l -
anything
everything
swam.
and
smaller
and
and
that
though it remains one of the lesser known mosasaurs from the lower
S m o k y H i l l C h a l k M e m b e r in Kansas, a sufficient number of specimens
now is available to provide a more accurate assessment of its size range in
comparison to T. proriger. N e w material shows that adults of this species
were significantly larger than the "one third" of the size of T. proriger
originally estimated by C o p e (1874) or even the specimens measured by
Russell (1967). In addition, large vertebrae from the lower one-third of the
S m o k y H i l l C h a l k M e m b e r (upper C o n i a c i a n ) indicate the presence of a
Tylosaurus that was about 89 m in body length. Although minor morphological features separate the two species, the fossil record indicates
clearly that by the end of the Coniacian (86 m y a ) T. nepaeolicus was approaching the same adult size observed in T. proriger remains from the lower
to m i d d l e Santonian and later.
T h e rivalry between C o p e and M a r s h involved mostly Eocene mammals
and dinosaurs, but their conflicts over mosasaurs were equally acrimonious. In
1868, C o p e traveled to Kansas and described several species of mosasaurs,
including Clidastes, Platecarpus, Mosasaurus, and one that he named Liodon, a
corruption of Owen's already existing Leiodon. Based on his preliminary examination of the mosasaur material, Cope developed some strange notions
about these reptiles, and in 1869, he classified them into a new order, the
Pythonomorpha, because he believed that they were snakelike in form. (In the
same paper, he erected the order Strcptosauria to include the elasmosaurs
with their vertebrae reversed.) Of the mosasaurs, he wrote (1869b):
We may now look upon the mosasaurs and their allies as a race of gigantic,
marine, serpent-like reptiles, with powers of s w i m m i n g and running, like
the m o d e r n Ophidia. A d d i n g a pair of short anterior paddles, they are not
badly represented by old Pontoppidan's figure of a sea serpent. T h a t terrestrial representatives, unknown to us, inhabited the forests and swamps of
the M e s o z o i c continents, and strove for mastery with the huge dinosaurs,
that also sought their shades, is p r o b a b l e . . . . T h u s in the mosasaurids, we
almost realize the fictions of snakc-like dragons and sea serpents, which
men have been ever prone to indulge. On account of the ophidian part of
their affinities, I have called this order the P y t h o n o m o r p h a .
Probably
inhabitant
shallow
an
oj
inshore
waters,
I 'lioplatccarpus
had the largest brain
oj any
mosasaur.
joined. If one found a particularly rich fossiliferous site, the other moved in
immediately, luring away the other's workers by offers of higher pay, destroying markers, and occasionally even stealing the o t h e r s specimens. T h e "bone
hunters" of C o p e and M a r s h went into the field a r m e d not only with picks
and hammers but with pistols and rifles as well. Of course, the guns were used
to hunt game and shoot buffaloes, and the various Indian tribes were more
than a little perturbed at being driven off their lands, but each man also felt
the need for protection against his rival's troops. (General George Custer and
his cavalry were slaughtered by S i t t i n g Bull and C r a z y Horse at the Little Big
H o r n in 1876, and a mere week later, the fossil hunter Charles Sternberg, who
was then working for Cope, was sent into the Black H i l l s of M o n t a n a to look
f o r dinosaur fossils.) Backed by the money of his uncle George Peabody ( w h o
had founded the m u s e u m at Yale and endowed a chair in paleontology f o t his
the vertebrae. Russell (1970) referred Ciidastes sternbergii to the genus Halisaurus,
so the current name of this species is Halisaurus sternbergi (Russell also respelled
the specific name, dropping the extra " i " ) . L i n g h a m - S o l i a r (1991b) didn't
agree with Russell that H. sternbergi should be referred to Halisaurus, and he
wanted to include Phospborosaurus (named by Dollo in 1889) in the genus
Halisaurus. H o l m e s and Sues (2000) argued that Phospbcrosaurus should probably remain distinct from Halisaurus.
Charles C a m p , who discovered the giant ichthvosaur graveyard in Nevada,
also found some new mosasaurs in California. In his 1942 publication " C a l ifornia M o s a s a u r s , " he wrote:
I he evolution of the mosasaurs from early Cretaceous varanoid lizards
such as Aigialosaurus is an instructional example of the changes that occur in
the passage from a land to a marine habitus in reptiles. T h e walking foot of
the land lizard became larger, with longer and more slender fingers in the
earliest known aquatic forms. Presumably these were webfootcd. S o m e of
the early mosasaurs were also slender-figured, with a loosely-knit, seaturtlc-like paddle. In advanced mosasaurs such as Mosasaurus and Kolposaurus the digits were adprcssed, the number of phalanges was increased,
and there was no doubt a thick, fibrous envelope around the hand, as in
whales.
C a m p created a new genus of California mosasaurs that he called Kolposaurus, which means "bay lizard" (from the Greek kolpos, meaning "bay" or
"gull " ) . He included two new species, Kolposaurus bennisoni (named for Allan
Bennison, the high school student who found the fossil in 1937) and K. tuckeri
( n a m e d for Professor W. M. Tucker, who found the second fossil). Both
specimens were large, long-tailed mosasaurs 30 to 40 feet long, bach had a
slender, pointed skull; a mouthful of sharp teeth; and very large eyes, which
suggest that this species was a fast-swimming predator that probably fed on
surface fishes. In 1951, having learned that the name Kolposaurus was "preoccupied" (by a nothosaur), C a m p changed the genus name to Plotosaurus, which
means " s w i m m e r lizard," so his California mosasaurs are now Plotosaurus
bennisoni and P. tuckeri.
At a m a x i m u m of 21 feet, the Platecarpus mosasaurs were exceptionally fast
and flexible, but their teeth were relatively small, suggesting a diet of small
fishes and cephalopods, especially bclemnites.* In his 1970 review of the
mosasaurs ol the Sclma Formation ( A l a b a m a ) , Russell wrote, "If bclemnites
normally formed a large proportion of the diet of the slender-toothed
plioplatecarpines
(Platecarpus,
Plioplatecarpus,
Ectenosaurus)
it might be expected
* Bclemnites were marine cephalopods similar to modern squids and cuttlefishes, in that
they were dartlike, rapid swimmers and could move tailward or tentacleward with equal
facility. (The word comes from the Greek belemnon, meaning "dart" or "javelin.") In those
rare fossils in which the soft tissue has been preserved, it can be seen that bclemnites had ten
tentacles of equal length, set with rows of little hooks instead of suckers, and, like the
modern cephalopods, they had an ink sac. The bclemnites' squidlike body enclosed a coneshaped internal shell (the phragmocone), which terminated at the tail end in a solid,
pointed element known as the rostrum or guard. The phragmocone resembled a straightened nautiloid shell, and the pro-ostracum corresponded to the calcified pen or gladius of
living squids and cuttlefishes, but no living cephalopod has a solid guard at the posterior
end, so the function of this bullet-shaped clement can only be guessed at.
seum of Natural History, they recognized it as having come from the M o o r e ville C h a l k Formation around Selma. It was a plioplatecarpine mosasaur that
somewhat resembled the N o r t h American genus Ectenosaurus and the African
Goronyosaurus, but it was different enough to warrant its own genus. T h e y
named it Selmasaurus russelli the generic name for the location, and the specific name for Dale Russell, "for his extensive work on the Mosasauridae."
Until quite recently, there was no evidence one way or the other to show
how mosasaurs were born. W i l l i s t o n (1914) wrote, " T h e legs were so c o m pletely adapted to an aquatic m o d e of living that the animals must have been
practically helpless on land, able perhaps to move about in a serpentine way
when accidentally stranded upon the beaches, but probably never seeking the
land voluntarily. . . . I f the mosasaurs were viviparous, a s were the ichthyosaurs, and probably the plesiosaurs, and as are some living land lizards,
the apparently entire absence of e m b r y o n i c bones associated with often nearly
complete skeletons of the mosasaurs is inexplicable; certainly some mosasaurs
must have died a short time before the birth of their young." In 1989, Russell
suggested that "perhaps, like some living reptiles . . . they sought the secluded
beaches of isolated islets and atolls in which to lay their eggs," but this seemed
unlikely to some, because these giant lizards would probably have been too
large and too ungainly to move themselves about on land to lay eggs.
But then Gorden Bell, one of the world's foremost authorities on mosasaurs, found the fragmentary remains of the mosasaur Plioplatecarpus primaevus
in South Dakota, along with the bones of two prenatal mosasaur embryos
(Bell et al. 1996). T h e bones were disarticulated, but this was attributed to the
scavenging by an opportunistic school of dogfish sharks (Squalicorax), whose
presence was evidenced by more than 2,000 shark teeth in the i m m e d i a t e
vicinity of the fossil embryos. It now appears that the mosasaurs, like the
ichthyosaurs, gave birth to live young at sea. ( T h e juvenile mosasaurs on Vega
Island suggested another behavior previously unsuspected: they might have
cared for their young as crocodilians d o . ) Further support for the idea of
viviparous mosasaurs came in 2001, when C a l d w e l l and Lee published a
description of a fossilized aigialosaur (Carsosaurus) with "at least four advanced embryos distributed along the posterior two-thirds of the trunk region. T h i s orientation suggests that they were born tail first (the nostrils
(which turned out to be s y n o n y m o u s with T. proriger). M o r e than threequarters of the material found by the Zangerl expedition belonged to the
species first found by Cope, Clidastes propython, which is also known from the
Niobrara fauna. In his reports, Zangerl noted that "the presence of turtles,
small pterodactyls, and toothed birds . . . indicated shallowing water and an
approaching shoreline" (Russell 1970), and it was on this basis that Russell
considered Clidastes a shallow-water mosasaur.
Shallow waters tend to be warmer than deeper waters, and the presence of
Clidastes, as well as Platecarpus and Globidens, lends "a definite Tethyan or tropical
cast to the mosasaur assemblage from the Mooreville C h a l k " (Russell 1970).
Tyrrell Museum of Alberta is currently excavating a fossil ichthyosaur whose skull was 18
feet long and whose total length has been estimated at 75 feet.
or nearly so. Dollo's original Hainosaurus bemardi came from Ciply, and there
are examples of five other mosasaur species Carinodens belgicus, Plioplatecarpus
houzeaui,
Prognathodon giganteus,
Prognathodon
solvayi,
and
an
unnamed
Halisaurus
species but most of the material has been referred to Mosasaurus umonnieri,
which Russell (1967) suggested s y n o n y m i z i n g with the American species
Mosasaurus conodon. But in 2000, L i n g h a m - S o l i a r restored Mosasaurus umonnieri
to the status of full species, writing that the "previous assignment to M.
conodon is rejected here." T h e original material was found at Ciply, through a
collaboration between Louis D o l l o of the Belgian Royal Institute and the
engineers L. Bernard and A. Lemonnier of the phosphate company. ( D o l l o
named Hainosaurus bemardi and Mosasaurus lemonnieri for these two gentlemen.)
T h e great majority of the material from the C i p l y phosphates belongs to the
species M. lemmonier, which is differentiated from the very similar but much
larger M. boffmanni by size and certain skeletal characteristics. T h e durophagous genera Globidens and Carinodens were also found in Belgium and the
Netherlands, demonstrating that these lowland countries represent some of
the most important sites in the world for the study of mosasaurs.
In 1998, Dutch paleontologist Rtitid Dortangs was fossicking in the cement q u a r r y at the Maastricht site when he noticed a bone protruding from
the limestone. It turned out to be a caudal vertebra of a large mosasaur, and
with his associates from the N a t u r a l H i s t o r y M u s e u m , D o r t a n g s began to
excavate the remainder of the skeleton. T h e y found an almost complete skull,
more tail vertebrae, the shoulder girdle, some teeth, and the rib cage. W h e n
they had removed and examined the bones, they "more or less routinely"
identified them as belonging to the giant Maastricht mosasaur M. boffmanni,
but "subsequent preparation revealed features and skull inconsistent with
such an assignment" ( D o r t a n g s et al. 2001). T h e y realized that this 45-foot
mosasaur was a new species of Prognathodon. An animal this size would have
had few natural enemies, but the bones were scratched as if from sharks' teeth.
It cannot be known whether the sharks killed the mosasaur and then fed on it
or came upon its carcass after it had died, but the scratches on the bones and
the sharks' teeth found alongside the fossil indicate that sharks had fed on the
mosasaur, which was named Prognathodon saturator. Saturator means "one who
gives satisfaction" in Latin, and the name was chosen because the mosasaur
Found in a quarry
in Maastricht, the
Netherlands,
Prognathodon
saturator is the
most recently
discovered mosasaur.
O u t s i d e of Kansas and the Netherlands, some of the best mosasaur material comes from Africa. In 1912, Robert Broom ( w h o would go on to discover
Australopithecus rohustus, now known as Paranthropus robustus) wrote about a frag-
At a length of ^
South Africa, that he felt was "manifestly not Tylosaurus proriger C o p e " but was
of the largest.
quite similar to
T. dyspelor, so he named it
Tylosaurus capensis. W h e n W. E.
and named the 25-foot-long Goronyosaurus nigeriensis* a new species that resembled no other mosasaur, and another specimen that resembled Halisaurus.
Thcagartcn L i n g h a m - S o l i a r has been working with the African mosasaurs
since 1988, when he published a paper ( u n d e r the name T. Soliar; L i n g h a m
was added later) on Goronyosaurus from the U p p e r Cretaceous of S o k o t o State,
Nigeria, which had been named for the Goronyo district by Azzaroli et al. in
1972. In a later discussion, L i n g h a m - S o l i a r (1999b) called Goronyosaurus "one
of the most enigmatic extinct marine reptiles, manifesting the largest number
of derived characters in the Mosasauridae." T h e skull of Goronyosaurus differs
from that of any other mosasaurs in that it is not tapered toward the front;
rather, it can be described as an elongated cylinder. It resembled a crocodile in
the way its teeth, including the first caniniforms in mosasaurs ( o n l y m a m m a l s
have true canines), fit into deep sockets in the opposing jaws. T h e estimated
length of an adult Goronyosaurus was 21 feet, m a k i n g it one of the larger known
mosasaurs. (For comparison, M. hoffmanni, the largest of the mosasaurs, was
more than twice as long.) T h e large teeth and powerful jaws suggest that it fed
on large fish and other reptiles. Its eyes were relatively small, but its sense of
smell was keen, and it probably fed much the way the marine crocodiles do: by
ambushing their prey and then tearing it apart with their powerful jaws and
teeth. L i n g h a m - S o l i a r (1999b) wrote:
In addition the long body form of mosasaurs, suited to ambush predation,
was a distinct advantage in the changing environment of the Late Cretaceous. T h e fast, highly evasive teleost fishes were experiencing a major
radiation at this time and the pursuit form of predatory tactics in marine
reptiles would have become energetically very expensive. It therefore seems
no coincidence that pursuit predators such as the ichthyosaurs became extinct at this point and the plesiosaurs were reduced from six families to two.
L i n g h a m - S o l i a r was so impressed with the teeth of Goronyosaurus that he
devoted two papers to them and their implications. In the paper quoted above
new information added here, vindicates the erection of a new genus, which can be tentatively assigned to the Tylosaurinac." A new subfamily might be warranted here, but so far,
(1999b), he showed the restored skull with the teeth in place, pointing out
Cretaceous of the
that the size and structure of the caniniform teeth "suggest that they were not
Coronyo district,
designed for impact against bone but rather shearing into flesh." He devoted a
Sokoto State,
Nigeria, the 21foot-long
Goronyosaurus
was one of the larger
later paper (2002b) only to the caniniform teeth and how Goronyosaurus might
have used them: "Taken together with skull strengthening features discussed
in a functional study of the skull of Goronyosaurus, the enlarged teeth . . . seem
to be a novel mosasaurian development of the dentition and a food trap. Such
mosasaurs. With
features arc more characteristic of some crocodiles and much earlier terrestrial
those of a crocodile,
this reptile was a
formidable predator.
swam in the Trans-Saharan Seaway, a part of the Tethys Sea extending from
the Gulf of Guinea in the west to the M e d i t e r r a n e a n in the northeast,
essentially bisecting the African continent. L i n g h a m - S o l i a r envisions Pluridens
"as an ambush predator, as are most mosasaurs, accelerating rapidly by means
of the long tail when prey was sighted."
"Recent studies," wrote L i n g h a m - S o l i a r (1994a), "show that mosasaurs
were also prevalent in Zaire and Angola. T h e picture therefore is that of an
almost continuous band of mosasaur localities stretching from Egypt, across
to M o r o c c o and Algeria, then southwards to Niger, Nigeria, Zaire, Angola,
and South Africa. Hence, African mosasaurs must today rival the historical
and present-day discoveries of two of the greatest mosasaur bearing regions
of the world, N o r t h America and B e l g i u m / T h e Netherlands." L i n g h a m Soliar found mosasaur fossils from A to Z Angola to Zaire in Africa. He
identified fragmentary mosasaur material mostly teeth referable to four
species in Zaire: Plioplatecarpus, Prognathodon, Halisaurus, and Mosasaurus. He examined a new species that had been found in Angola and named Angolasaurus
bocagei, but "reassessment of the material indicates that it clearly belongs to a
new species of Platecarpus," geographically the most widespread member of the
Mosasauridac and inhabiting the waters of ( w h a t is n o w ) N o r t h America,
Europe, Africa, and N e w Zealand. Of Plioplatecarpus marshi, he wrote, " W i t h
needle-sharp, strongly backwardly recurved teeth, it drew in prey by what is
known as ratchet feeding similar to that of snakes 'walking' the jaws over the
p r e y . . . . It is interesting that the animal also shows early signs in its m o r p h o l ogy of adopting a penguin-like m o d e of locomotion, underwater flight, that
would have been highly useful in the complex, crowded habitats it is believed
to have inhabited."
Like sub-Saharan Africa or Kansas, for that matter the Ncgev Desert
of Israel is one of the last places one w o u l d expect to find the remains of
extinct aquatic reptiles. But in Cretaceous times, these areas were all underwater, and the waters were occupied by mosasaurs. In 1993, workers at the Oron
phosphate field 30 miles south of Be'cr Sheva unearthed a fossil mosasaur, and
it was shipped to Copenhagen, where preparator Stcn Jakobsen worked on it
for two years. Paleontologists N i e l s Bonde and Per Christiansen then began
the scientific analysis of the 5'/2-foot-long skull that was encased in silicified
sandstone as hard as concrete. T h e wide, heavy skull, with its massive jaw
teeth and especially large, curved palatal teeth, suggested that Oronosaurus
nicknamed for the place it was found was a mega-predator, well designed to
prey on large vertebrates. From the length of the skull, Bonde and C h r i s tiansen estimated the total length of the animal at about 40 feet, making
Israel's first mosasaur one of the largest known. T h e skull, which is the largest
fossil of anything ever unearthed in Israel, now resides at the Geological
Institute of the University of Copenhagen, but because it belongs to Ben
Gurion University in Be'er Sheva, it will be returned there when a museum is
built in which to display it.*
In September 2002, the description was finally published in the Journal of
Vertebrate Paleontology.
Christiansen
and
Bonde
named
the
new species
Prog-
Tylosaurus baumuriensis,
Mosasaurus mokoroa,
and
new species
she
to Rikkisaurus Wiffen,
overtoni, and
MosasaurusJlemingi arc reassigned to Moanasaurus. Phylogcnctic analysis indicates that the moanasaurines are sister-group to the Plotosaurini (Mosasaurus and Clobidens); clidastines arc the sister-group to moanasaurines and
Plotosaurini. M o s a s a u r fossils occur in giant concretions formed within
near-shore marine elastics, the beds of which are marked by intense bioturbation. N e w Zealand mosasaurs were occupying nearshore environments
and may have used coastlines to radiate and migrate between continents.
mosasaur that
crushed shells with
its round teeth, hence
fast-moving prey items and snagging them with their big, sharp teeth. T h e
genus known as Globidens ("round teeth"), first discovered by Gilmore in
A l a b a m a in 1912 and subsequently unearthed in Kansas and South Dakota,
had rounded teeth instead of the more typical conical spikes. From the partial
jawbones that have been excavated, it has been estimated that Globidens reached
a length of 20 feet, and perhaps even more. Although a reptile of this size
Alabama, Belgium,
and the Netherlands.
would have been a formidable predator, its rounded teeth were designed to
crush shellfish probably ammonites. Because we know that mosasaurs occasionally fed on ammonites, it is not that much of a leap to imagine a mosasaur
species that evolved to eat only these cephalopods. (A modern analogue is the
horn shark, Heterodontus sp., with pavement-like teeth in the back of its jaws
that are used specifically for crushing the hard shells of oysters and clams.)
Two tooth types arc identified in Globidens: a spherical type, fairly smooth and
with a small nubbin or point, and a subspherical form, with a highly striated
surface and a large apical nubbin.
Smaller than Globidens but with similarly rounded teeth were the mosasaurs
Carinodens belgicus and Cjraasi, which were only 10 to 12 feet long and probably
searched the sea floor for brittle m o l l u s k s and sea urchins to crack open.
(Carinodens was originally named Compressidens by Dollo in 1924.) Globidens and
Carinodens from the U p p e r Cretaceous of Belgium and the Netherlands were
the first marine reptiles since the placodonts of the Triassic and the ichthyosaur Grippia from the Triassic of Spitsbergen that were specialized for
feeding on shelled animals. Based on just a few teeth on a slender jaw,
Lingham-Soliar (1999a) reconstructed the entire dental row of Carinodens to
show, in lateral view, pointed teeth anteriorly, triangular teeth toward the
middle, and rectangular teeth posteriorly. Both the tooth shape a n d the
wrinkled surface played an important part in the stresses the teeth were
subjected to. Carinodens probably fed on thin-shelled invertebrates and Globidens on thicker-shelled invertebrates and vertebrates. T h e round-toothed
mosasaurs were the only ones that did not have teeth on the pterygoid bones
of the palate; crushed shellfish did not have to be "walked" to the throat like
struggling vertebrates. Feeding on hard-shelled animals ( d u r o p h a g y ) also
occurred in other mosasaurs such as Mosasaurus boffmanni, Prognathodon, and the
tylosaurs, although in these forms it was probably opportunistic and part of a
wider feeding strategy.
Louis Dollo (1904) was among the first to record the feeding potential of
mosasaurs when he showed the fossilized remains of a large turtle in the gut
cavity of the 50-foot-long mosasaur Hainosaurus found in Belgium. Even with
its hinged lower jaw, it remains a mystery how Hainosaurus could have swallowed the t u r t l e s carapace ( L i n g h a m - S o l i a r 1992b). Said by some to have
been the largest of all mosasaurs, Hainosaurus has also been found in the
Kristianstad Basin in S k a n e , southern Sweden, and ( m a y b e ) in the Pierre
S h a l e of M a n i t o b a . T h e M a n i t o b a specimen, found in the pits of the Pembina M i n i n g Company, was named Hainosaurus pembinensis and was described
in 1988 by Betsy N i c h o l l s , now of the Royal Tyrrell M u s e u m . It was the first
record of Hainosaurus from N o r t h America.* T h i s species is the longest of the
mosasaurs because it had more precaudal vertebrae than any other species; it
had 53, compared with the 35 of Tylosaurus. T h e teeth are minutely serrated,
and the narrow skull is shaped like an arrowhead. Although unspecialized
feeders, these giant mosasaurs were highly specialized in their killing mechanism, involving not only a huge and somehow expandable mouth but also a
large, solid rostrum at the tip of the snout that could be used as a battering
ram to stun or kill prey by smashing into it, not u n l i k e the way a bottlcnose
d o l p h i n uses its "beak" to fight off sharks.
An animal with a built-in battering ram is likely to use it, and there is
evidence that one large mosasaur killed another by a powerful blow to the
head. T h e victim was a subadult Mosasaurus hoffmanni, and the suspected perpetrator was Hainosaurus bemardi. Examining a cast of the braincase of M. hoffmanni in the collection of the Institut Royal des Sciences Naturelles de
Belgique, L i n g h a m - S o l i a r (1998c) found "some unusual damage," consisting
of a severed and displaced cerebellum, that could only have come from "a
powerful concentrated blow to the prootic region of the braincase. T h e
prootic would have sprung inwards, facilitated by ligamentous sutures, bro-
2000b),
the Institute Royal in Belgium, in both cranial and postcranial measurements there are
considerable intraspecific variations. I personally would nor have created this new North
American species, with all its ramifications, without a thorough investigation of the bountiful Tylosaurus specimens in the USA. In Dollo's day I think it was understandable, particularly with his penchant for telegraphic descriptions."
shown crushing an
ammonite whose ink
sac has hurst. The
head of the mosasaur
was modeled after
a drawing by
Lingham-Soliar that
appeared on the
ken internally thereby causing the brain to be severed and sprung out again."
W h a t sort of animal could ( o r w o u l d ) ram a gigantic mosasaur? Probably an
animal that was designed to do just that. T h e tylosaur Hainosaurus had a large,
bony rostrum; was capable of bursts of speed; and had a mouthful of strong,
sharp teeth and an e n o r m o u s appetite. Of course, we will never know what
caused the death of M. hoffmanni, but the clues, as read by "Sherlock" L i n g h a m Soliar, point strongly in the direction of Hainosaurus. It had the weapon, the
opportunity, and the motive.
In 1991, L i n g h a m - S o l i a r wrote a paper called " L o c o m o t i o n in M o s a s a u r s "
in which he reiewcd the early speculations on mosasaur s w i m m i n g , many of
which had the reptiles undulating their entire bodies like snakes or eels. After
a careful analysis of the structure of the vertebrae, he concluded that they
cover of the
European magazine
Science Spectra
in
1999.
maneuverability than the deeper-water types, "and it seems that the unusual
structure of Plioplatecarpus marsbi led to adaptations for pursuit of fast a n d /
or elusive prey in a structurally complex environment."
T h e idea of underwater flight in mosasaurs has led to a spirited controversy within the palcontological community. For instance, Nicholls and Godfrey (1994) accepted Lingham-Soliar's original interpretation, but not the
revised version. T h e y recognized that the spine of Plioplatecarpus marsbi was
stiffened by especially heavy vertebrae, but they said that in all other subaqueous "fliers" (such as p e n g u i n s ) the tail is greatly reduced, and "mosasaurs
traditionally have long, laterally compressed tails, and there is no reason to
believe that Plioplatecarpus was any different in this respect." T h e y contended
that the huge flippers of this mosasaur would not have been particularly
effective as wings, and that the powerful pectoral girdle "may be associated
with a number of other functions, such as an increase in maneuverability in a
structurally complex environment." Furthermore, "sharks which shake their
prey have well-developed pectoral girdles. Sharp movements of the pectoral
fins are transmitted to the head to achieve the vigorous shaking needed to
dismember the prey. T h e massive deltopectoral crest and pectoral girdle seen
in Plioplatecarpus might have allowed them to feed in this fashion." In his
discussion of "Carrier's Constraint," Cowen (1996) wrote that the body was
indeed stiffened, but not for underwater flying. W h y was it stiffened? "I
suggest," wrote Cowen, "that Plioplatecarpus was beginning to solve Carrier's
Constraint, by decoupling flexure of the thorax from s w i m m i n g propulsion,
which increasingly involved only the posterior of the animal. Such an adaptation would only be important for an animal s w i m m i n g at sustained speed in
surface waters. Therefore, I suggest, this animal was the best sustained surface
swimmer among mosasaurs, even if it d i d not fly underwater."
If the locomotion of Plioplatecarpus marsbi "increasingly involved only the
posterior of the animal," one would expect a tail fin at least as prominent as
those of Ciidastes, Plotosaurus, and the various Mosasaurus species, but the neural
spins of the caudal vertebrae of Plioplatecarpus marsbi are the shortest known in
mosasaurs, indicating the unlikelihood of a reasonably developed tail fin.
Surface s w i m m i n g in a large mosasaur such as P. marsbi would be very expensive energetically because of increased drag resulting from wave action and
water movement for several feet below the surface. In the 1999 paper in which
they described a plioplatecarpine mosasaur from an estuarine environment,
H o l m e s , Caldwell, and C u m b a a rejected Lingham-Soliar's reconstruction and
biomechanical analysis of P. marshi. T h e y wrote that "his analysis required
extensive reconstruction of l i m b elements, most of which were not present in
the specimen available to him." L i n g h a m - S o l i a r hypothesized that the paddle
of P. marshi was used in subaqueous flying, but H o l m e s et al. believe that
Plioplatecarpus did not "fly" at all but paddled for initial acceleration with its
foreflippers, then tucked them in (presumably to reduce d r a g ) and used its tail
for sustained swimming. T h e foreflippers would also be used to effect sudden
changes in direction, both laterally and vertically.
In the past, paleontologists often examined and analyzed fossils, believing
that their responsibility began and ended with ostcological descriptions (e.g.,
"the tubular bassioccipitalia are partially concealed by the undcrlap of the
p t e r y g o i d s " ) , but nowadays, most paleontologists believe that they can and
should postulate a lifestyle from the fossil evidence. L i n g h a m - S o l i a r is a
charter m e m b e r of the m o d e r n group. In a 1993 article entitled " T h e M o s a saur Leiodon Bares Its Teeth," he indulged in the usual descriptions of the bones
(which arc, in this case, mostly the teeth of the t i t l e ) , as in, "anteriorly there is
a small rounded edentulous process," but his analysis was directed toward an
understanding of how the mosasaur might have used those teeth. After differentiating Leiodon ("smooth teeth") from other mosasaur species (the name
was first employed by Richard Owen in a series of papers in 18401845) and
laying the requisite ostcological and taxonomic groundwork, he gets to the
core of his discussion:
evidently tried to swallow the entire ammonite, pulling it as far back into
the throat as possible.
T h i s is what happened to the ammonite:
T h e shell walls bear numerous subround perforations, crushed areas, and
dents of several sizes which were made by the teeth of a mosasaur. T h e s e
m a r k s are present on both sides of the shell, but arc best developed on the
left side, which shows an almost complete set of maxillary impressions. A
dorsal sector of the conch has been fractured and slightly displaced, and the
living chamber, which makes up about half of the outer whorl of Placenticeras,
has been severely crushed and slightly torn at its apertural margin.
T h e r e are those, however, who believe that the d r a m a t i c event, presented in
such g r a p h i c detail by Kauffman and Kesling, d i d not happen at all. In his
1998 book Time Machine, cephalopod specialist Peter W a r d devotes an entire
chapter to " T h e Bite of a Mosasaur," in which he recounts his observations of
a sea turtle attack on a nautilus in captivity in N e w Caledonia, during which
the nautilus's shell was fragmented like a porcelain plate hit with a hammer.
T h e n he tells of his examination of a collection of Placenticeras shells in which
the holes "seemed to conform to the sizes and shapes produced by limpets."
He includes a discussion of graduate student Erica Roux's attempt to make
round holes in nautilus shells, a close approximation of unfossilized a m monite shells:
Erica constructed an artificial mosasaur jaw. It did not look much like the
real tiling, being fabricated of metal with series of teeth made out of nails
and screws, but nevertheless it approximated the real thing in many ways.
T h e "teeth" descended onto the shell surface just as a mosasaur jaw would
have, and a gauge attached to the jaw showed the amount of pressure
needed to produce a break. A nautilus shell was put between the jaws and
the type of damage inflicted on the shell was observed. For several days
[the l a b ] reverberated with cracks and snaps, as shell after shell fell victim
to those jaws of death. An a r m y of mosasaurs could not have had so much
fun. A n d in all the carnage that ensued, not once was a round circular hole
approximating the size of a m o s a s a u r s tooth ever produced.
Like many others, Ward believes that mosasaurs did indeed eat ammonites;
they just crushed the shells to get at the soft, edible bits.
In 1998, Kase et al. published their findings in "Alleged M o s a s a u r Bites on
Late Cretaceous Ammonites Are L i m p e t (Patellogastropod) H o m e Scars."
Using a robot mosasaur jaw, they demonstrated that the pressure exerted by a
mosasaurs teeth would utterly fracture a cephalopod's shell (there being no
fresh ammonites available, they used nautiluses). M o r e significantly, they
examined the holes supposedly m a d e by the teeth ol mosasaurs on a m m o n i t e
shells and found clear evidence of limpets "grazing" on the shells with their
radular teeth. T h e y concluded:
We do not deny that mosasaurs may have preyed on ammonites by crushing. Any that escaped swallowing had the potential to be preserved as
fossils. However, it is difficult to determine whether any resulting fragments are attributable to mosasaur bites or to other geologic processes. We
cannot accept that both limpet home scars and tooth punctures are present
on the ammonite shells. It is improbable that two unrelated k i n d s of
punctures should always occur at the same localities and only in a narrow
stratigraphic interval. We conclude that all the holes in ammonites attributed to mosasaur bites are l i m p e t hole scars that were altered by
diagenesis [the chemical, physical, and biological changes undergone by a
sediment after its initial deposition, and during and after its lithification,
exclusive of surface a l t e r a t i o n ] . O u r findings are important in revising Late
Cretaceous marine food webs and the alleged role of mosasaurs as a m monite predators.
Also in 1998, Seilacher wrote " M o s a s a u r , Limpets or Diagenesis," in which
he said, " T h e claimed 'mosasaur bites' are probably all caused by limpets
rasping on necroplanktonic Placenticeras shells, compactual puncturing of the
pits and diagnostic beveling of the rims." In other words, Seilacher believed
that the ammonites were already dead when they were colonized by the
limpets.
But Tsujita and Wcstcrmann (2001) believe that even if limpets made some
of the holes, it is highly unlikely that they m a d e all of them, and they support
Kauffman and Kcsling's parsimonious suggestion that h u n g r y mosasaurs
punched holes in the a m m o n i t e shells with their powerful jaws and teeth.
Tsujita and W e s t e r m a n n examined more than 150 specimens of Placcnliceras
from the U p p e r Cretaceous Bcarpaw Formation of southern Alberta and
found, as had Kauffman and Kesling, that many of the shells exhibited holes
in straight rows or even in a V-shaped pattern, which conformed to the jaw
shape of mosasaurs but was highly unlikely to result from random assemblages of limpets. Furthermore, they said, "fossil limpet shells arc much too
rare in the Bcarpaw Formation to account for all the perforations." T h e
designers of the robot jaws employed on nautilus shells assumed "that
mosasaurs always closed their jaws in a violent snapping motion" that shattered the shells, but that assumption is unwarranted; indeed, the "unusually
loose jaw structure of mosasaurs probably allowed for a great deal of controlled variation in biting habit," and they would therefore not crush the
shells as the robots did. " M a n y aspects of the perforations and associated
features preserved in Placcnticcras shells that can easily be accounted for by the
mosasaur-bite hypothesis, are impossible to explain by the limpet hypothesis. . . . We confidently conclude that the vast majority of perforated specimens of Placenlkeras, at least those from Alberta, record evidence of prcdation
and that the mosasaur-bite mark hypothesis has been unjustly dismissed by
proponents of the limpet home scar hypothesis."
I lean toward the mosasaur-bite hypothesis, and I believe that Kauffman
and Kesling are correct. Tsujita and W e s t e r m a n n arc convincing, in that
limpets were unlikely to have arranged themselves in patterns that so closely
resembled the tooth rows of mosasaurs, and their detailed analysis of the jaws
of mosasaurs explains why a m m o n i t e shells would not be shattered. It is
unlikely that Kase, Seilacher, Ward, and the rest of the proponents of the
limpet scar hypothesis are going to concede after all, Tsujita and Westermann reexamined the same evidence they did, not something new and
definitive and the argument is certainly not settled. To understand the
mosasaurs' feeding habits, we really need Peter Ward's time machine.
Like the other marine reptiles, the mosasaurs were probably partially
endothcrmic, but the question of where the energy came from for sustained
locomotion is not often addressed ( L i n g h a m - S o l i a r does not mention i t ) .
Massarc (1994) wrote, " T h e physiology of the animal is the most important
factor in estimating sustained s w i m m i n g speeds. Mosasaurs, related to m o n i tor lizards, were almost certainly cctotherms." ( M o n i t o r lizards, believed by
some to be closely related to mosasaurs, are surely ectotherms.) Although he
d i d not know il they dived deeply, Russell (1967) indicated that Tylosaurus and
Platecarpus frequented the deeper parts of the water column on a regular basis.
Although the surface waters of the interior seaway might have been w a r m e r
than the ocean, the depths were certainly cool, and we are pretty sure that
some mosasaurs were capable of deep dives (estimates of the greatest depth of
the interior seaway rarely exceed 600 feet).
W h e n they examined the fossilized vertebrae of some N o r t h American
mosasaurs, Rothschild and M a r t i n (1987) found evidence of avascular necrosis in two of the most c o m m o n genera, Tylosaurus and Platecarpus. T h i s bone
disease, which was present in nearly every skeleton they examined, occurs
when the blood supply to the bones is cut off, and it indicates an episode of
decompression sickness ("the b e n d s " ) , which results from nitrogen bubbles
entering the bloodstream under pressure as an animal ascends after a deep
dive. No such diseased vertebrae were found in specimens of Clidastes, which
was not a particularly deep diver, as far as we know. Necrotic bone in
Tylosaurus and Platecarpus suggests that these were deep-water species that m a y
have dived too deeply and too often.
W i l l i s t o n (1898), the earliest authority on mosasaurs, believed that Clidastes
was a swift surface hunter, but given what we know about living ocean
predators, such as dolphins, sprinting in pursuit of prey at the surface is a very
uncommon technique. Dolphins play or bow-ride at or near the surface but
do most of their hunting while submerged. T h e same is true of sharks, which
may take their prey at the surface but stalk it from below. ( T h e great white
shark, which feeds on seals and sea lions, approaches them from below and
consummates the attack at or near the surface.) T h e fish variously known as
the dorado, mahi-mahi, or Coryphaeua hippurus is probably the closest thing to a
"swift surface hunter." Considered one of the fastest of all fishes, Coryphaeua
chases flying fishes at the surface and occasionally even catches them in the air.
Like dolphins and unlike dorados and sharks mosasaurs had to surface to
breathe, so the opportunity to take prey at the surface was always there.
W i l l i s t o n believed that Tylosaurus was a predator of other marine reptiles and
that Platecarpus was a deep diver. M a r t i n and Rothschild (1989) also wrote that
in the Pierre S h a l e of S o u t h D a k o t a they "have collected skeletons of these
giant forms [ o f m o s a s a u r s ] in the same area as the remains of giant extinct
squids that m a y have been 6.2 to 9.2 meters [20 to 30 feet] long. It is possible
that Tylosaurus m a y have dived to great depths to capture squid, as the modern
s p e r m whale does now."
On the deep-diving ability of Platecarpus, A m y S h e l d o n (1997) disagrees
with M a r t i n and Rothschild. S h e wrote, "Platecarpus shows pachyostosis [an
increase in bone d e n s i t y ] which requires an increase in lung volume. T h i s
suggests a shallower and narrower range of neutral buoyancy." Sheldon believes that "bone microstructure seems to correlate with ecology"; that is,
those a n i m a l s with increased bone density are negatively buoyant even in
shallow water, and the heavy bones, like a diver's weight belt, keep the animals
submerged. ( S i r e n i a n s , with some of the densest bones of any animals, inhabit
only shallow waters, and Steller's sea cow [Hydrodamalis] probably could not
submerge at a l l . ) S h e l d o n wrote, " M a n y cetaceans, such as dolphins and some
whales, ichthyosaurs, and some turtles, have very porous, light bone, and
many s w i m swiftly."*
An inclination to get the bends is what evolutionary biologists call "mal* In one living whale species, the bones are the densest known for any animal. Blainville's
beaked whale, also known as the dense-beaked whale (Mesoplodon iensirostris), is a species
known from stranded specimens and occasional sightings at sea. Examining the rostrum
(the forward, pointed portion of the upper jaw) of a specimen in the Museum National
d'Histoirc Naturelle de Paris, Buffrcnil et al. ( 2 0 0 0 ) found that the bone was 2 2 percent
denser than any other known mammalian bone. Since no one has ever seen a dense-beaked
whale diving or hunting, the authors could only speculate as to the function of this
incredibly heavy bone. They discounted the idea that it might be used for intraspccific
fighting because its density renders it brittle, and "bones adapted for shock loading, such as
deer antlers, have the opposite structural characteristics." It might, they thought, be used as
an ultrasound transmitter, but not enough information is known on sound production in
beaked whales. Does it help in deep diving? Probably not. T h e authors concluded: "In the
absence of experimental investigation, the true functional role of this feature is largely a
matter of conjecture." II we can't ligurc om why .1 living whale needs such ,1 dense rostrum,
imagine how difficult it is to understand the physiological requirements of creatures that
have been extinct for 1 0 0 million years.
adaptive" a development that should produce a population that will eventually be doomed by its own habits. ( H u m a n s who get this disease are
operating lar outside the regular parameters of their b i o l o g y . ) D i d such a
situation contribute to the downfall of the mosasaurs or at least those that,
like Tylosaurus, were believed to be the deepest divers? M a r t i n and Rothschild
(1989) do not think so:
It seems unreasonable to assume that every deep dive would result in
decompression syndrome. If this were the case, how could deep diving
behavior have evolved? It seems likely that decompression events were the
direct result of crises such as the need to escape from predators or injudicious pursuit of p r e y . . . . D i d decompression syndrome have any role in
the final extinction of the mosasaurs? We do not think so. T h e fact that
they lasted for at least 25 million years a long time by most standards
suggests they were relatively successful organisms. . . . T h a t the final
disappearance of the mosasaurs occurred simultaneously with the extinction of such dissimilar organisms as dinosaurs and various types of phytoplankton strongly suggests that the cause was something other than a
special maladaptation.
For the past 30 years, M i k e Everhart has been uncovering fossil reptiles
from the Kansas shales and has found numerous mosasaur bones that show
distinct evidence of shark attack. From teeth embedded in the m o s a s a u r s
bones, the species of shark can be identified: it was Cretoxyrhina mantelli, a
lamnid that is known to have reached a length of 20 feet. After excavating
most of a fossilized mosasaur, Everhart realized that the ribs on the reptile's
right side had been bitten completely through. We will never know whether
the shark attacked a living mosasaur or scavenged a floating carcass. ( S i n c e the
extinct shark had no common name, Everhart christened it ginsu shark, "because it fed by slicing up its victim into bite-sized pieces." It is now popularly
known as the ginsu mako.) Cretoxyrhina is known from the fossil faunas of
Africa, Europe, and N o r t h America, and C. mantelli, described by Agassiz in
1843, is common in the U p p e r Cretaceous sediments of the Western Interior
Seaway, the warm, shallow sea that inundated central N o r t h America during
the Cretaceous. From the shape of the teeth, it is clear that Cretoxyrhina was
long, with a laterally compressed tail with the tip pointed downward and a
skull that was nearly identical to that of the mosasaurs. Both had a prominent
intramandibular joint, which opened the lower jaw unusually wide to help in
the bolting of large food items, suggesting an affinity of the aigialosaurs and
mosasaurs with snakes.
T h e sudden disappearance of these powerful, effective predators is, according to L i n g h a m - S o l i a r (1999a), "an enigma of the K - T extinction." He wrote,
" T h e i r feeding potential, aided by an arsenal of tools crushers, gougcrs,
The
mosasaur
Tylosaurus
attacking
mosasaur.
smaller
Ranging
in size from 10 to
JO feet, the tylosaurs
were the major
predators of the
Niobrara
the
Sea
Cretaceous
period.
during
But if, as M i c h a e l Lee (1997) has written, snakes and mosasaurs are sister
groups (both arising from a single c o m m o n ancestor), the mosasaurs are not
officially extinct at all. Lee combined mosasaurs and snakes in a group he
named Pythonomorpha, a name originally coined by C o p e in 1869 to include
the mosasaurs and snakes and based on the mythological serpent Python,
which was produced from the m u d left after Deucalion's flood. M c N a m a r a
and Long (1998) summed up Lee's unanticipated analysis by writing:
True terrestrial snakes appear by the Late Cretaceous, at the same time that
the radiation of mosasaurs was peaking. For many years the origin of
snakes was clouded in mystery due to lack of adequate fossil evidence. But
now Lee and Caldwell have blown away the cobwebs, by taking a fresh,
rigorous, and very detailed look at the known m a t e r i a l . . . . So when you are
next out snorkeling and are startled by a sea snake [ M c N a m a r a and Long
are Australians] it may not only be some highly derived snake that you are
frantically paddling away from, but all that remains of a great radiation of
a great tradition of aquatic reptiles that once dominated the seas.
In 1966, in response to the confused and inconclusive state of the study of
snake evolution at that time, Alfred S h e r w o o d Romer wrote, " i n contrast to
the extinction or seeming evolutionary stagnation of other reptile types, the
snakes are today a group of reptiles still 'on the m a k e . ' " M i n t o n and H e a t wole (1978) wrote, " T h e snakes appeared late in the M e s o z o i c , and there is
some evidence that, quite early in their history, they produced some huge
marine species. Apparently these giant sea snakes were not very successful, for
they endured but a short time and left a very scanty fossil record." According
to Heatwole (1999), "Other snakes from the Cretaceous are known only from
incomplete fossils and as no good skull material exists the most that can be
said of them is that they were snakes with some characteristics intermediate
between those of lizards and modern snakes."
T h e earliest snakes are known from isolated and rather uninformative
vertebrae. T h e best preserved early snakes had an elongate body, reduced
limbs, and adaptations for chemosensory hunting. T h e y are descended from
lizards and manifest many lizard-like characters. In 1997, Caldwell and Lee
published a description of Packyrhachis problematicus, a fossil snake that was
evidence was not compelling, and later workers have been reluctant to accept
this view." In their r e v a l u a t i o n , Lee and his colleagues believed that they
showed that Pacbyopbis was indeed a very primitive snake. T h e fossil was found
in East Herzegovina, in the same locality as another snake known as Mesopbis
nopscai, which is now lost. T h e fossil of Pacbyopbis is smaller than that of
Pacbyrbachis, but its ribs are much heavier, indicating that it was not a juvenile
of the latter form. T h e thickened bone a condition known as pacbyostosis is
characteristic of marine animals because it increases their density, which
strongly suggests that this species ( a n d also Pacbyrbachis) were marine. " H o w ever," wrote the authors, "at the moment, whether or not all snakes went
through a marine phase in their evolution remains equivocal."
In 2001, Caldwell and Albino published a discussion of the paleoenvironment and paleoecology of the marine snakes Pacbyopbis and Pacbyrbachis and the
terrestrial snake Dinilysia. As mentioned earlier (Scanlon et al. 1999), Pacbyrbachis was shown to have inhabited interreef systems of the Tethyan seaway,
and with its small head and muscle attachments that correspond to those of
modern striking snakes, it was probably competent at plucking its prey from
within cracks and crevices. Because the type specimen of Pacbyrbachis contains a
partial tooth plate from a pyenodont fish, it might have eaten fairly large prey
and been able to spread its jaws widely, as do many m o d e r n snakes. At a
length of 4 feet, Pacbyrbachis was nearly twice as large as Pacbyopbis, but because
they shared many physical characteristics, their hunting methods were probably the same.
T h e descent of snakes is one of the most contentious areas in vertebrate
biology. Tchernov et al. (2000) were more than a little critical of the conclusions of Lee et al. (1999) and wrote, "Haasiophis and Pacbyrbachis have no
particular bearing on snake-mosasaurid relationships or snake origins. . . .
Basal snakes, including basal macrostomatans, retain rudimentary hind limbs,
which, however, remain much more incomplete than those of Haasiophis." But
as Caldwell (personal communication 2002) notes, he and Lee "never said
that they did [have such a b e a r i n g ] . It is also very important to realize that
Tchernov et al. did not repeat the study of Caldwell and Lee (1997) which
included all squamates and a number of fossil lizards, but rather restricted
their analysis to only snakes. T h e y rearranged the ingroup relationships of
palatal
Snakes have developed a rigid skull structure from which the highly flexible
jaws are suspended, which facilitates the engulfment of large prey items in
some cases, larger than the snake's head. In 1999, with Gorden Bell and M i chael Caldwell, he wrote, "Here we present evidence that mosasaurs large,
extinct marine lizards related to snakes represent a crucial intermediate
stage. Mosasaurs, uniquely among lizards, possessed long, snakelike palatal
teeth for holding prey. Also, although they retained the rigid upper jaws typical of lizards, they possessed highly flexible lower jaws that were not only
morphologically similar to those of snakes, but also functionally similar
In
terms of skull structure, the large, limbed marine mosasaurs were functionally,
as well as phylogenetically, intermediate between the lizards and snakes."
In a 2000 article delightfully entitled "Nice Snake, S h a m e about the Legs,"
Michael Coates and M a r c c l l o Ruta reviewed the hypotheses about the evolutionary origins of snakes. T h e y wrote:
Packyrhachis prohlematicus from Israel rapidly assumed a central position in the
debates about snake phylogcny. It has miniature hindlimbs articulated with
a rudimentary pelvic girdle, but sadly, its feet arc missing. Currently described from only two specimens, it was originally interpreted as an aquatic
relative of terrestrial varanoids and was therefore thought to be morphologically convergent with snakes. However, it has recently been interpreted as a true snake, based on its loosely articulated upper jaws, intramandibular joint, the condition of several skull bones, the absence of
forelimb and pectoral girdle, body elongation and vertebral structure.
T h e y said that according to Caldwell and Lee, Pachyrhachis is the most primitive known snake and "has gained classic 'transitional' status, bridging the gulf
between two highly specialized squamate clades," the mosasauroid-snakc
shared ancestry and modern snakes. R i e p p e l and Zaher take an opposite view,
maintaining that the scolccophidians (extant blind snakes) are the most
primitive snakes. T h e questions of snake developmental evolution are not
resolved. Coates and R u t a concluded: "it is worth remembering that phylogcny is an ongoing research program and that large parts of the evolutionary
tree remain unwritten, unexplored, and deeply uncertain.
As if to show that the derivation of snakes from lizards isn't so strange after
all, W i c n s and Slingluff (2001) published " H o w Lizards Turn into Snakes: A
Phylogcnetic Analysis of Body-Form Evolution in A n g u i d Lizards." For the
most part, anguid lizards already look like snakes: they arc the "glass snakes"
(Ophisaurus, confusingly translated as "snake l i z a r d " ) and " s l o w w o r m s " (Anguis
jragilis) of Africa, Europe, and Asia, which have no legs at all; and the alligator
lizards (Gerrhonotus) of western N o r t h America, which have extremely short
legs and long, sinuous bodies. To the layperson, it is obvious that they are not
snakes because they have closable eyelids and a notched (as opposed to a
forked) tongue. T h e y can shed their tails to escape predators, something no
snake can do. ( T h e glass snake is so named because besides being able to shed
its tail, it can shatter it into several pieces to distract predators even more.)
Any study labeled "A Phylogenetic A n a l y s i s " is going to be heavy going for the
layperson, and this one is no exception. It is replete with sentences like this,
selected at random: " T h e best-fitting model was then used in a heuristic
search to find the overall best likelihood t o p o l o g y using trce-bisectionreconnection branch swapping and 10 r a n d o m addition sequence replicates."
If you're not a phylogcneticist, such a sentence probably won't help you
understand how lizards turn into snakes, but the authors summarized their
findings by writing, " O u r results support the hypothesis that limb reduction
is correlated with body elongation and that digit loss is correlated with limb
reduction."
In the chapter " W o n d e r of the Kansas P l a i n s " in his 1887 book Sea and Ixmd:
An Illustrated History of the Wonderful and Curious Things of Nature Existing before and
since the Deluge, J. W. Bucl expressed the public's amazement at the discovery of
gigantic sea lizards in the b a d l a n d s of the American West:
T h e fabulous monsters that were believed in in the olden times, the
dragons, serpents, etc., are thrown in the shade by these truly ancient
monsters that once swam in the ocean that finally became land-locked, and
the bottom of which is now raised high above the water level. T h e shore
line of the ancient ocean is distinctly marked. Imagine the water between
N e w York and L o n d o n a dry plain, its whales and fishes stranded in the
mud, on the sides of the great hills, and on the plateaus that we know exist,
an idea can be formed o f the tnauvaise terres. Professor M a r s h says that in
one place he counted from his horse the r e m a i n s of five huge monsters
spread upon the plain. One of the largest of these, a reptile called the
l.iodon, exceeded in size the largest whale.
From the moment that the grand animate de Maastricht was hauled up from the
limestone m i n e in 1780, mosasaurs have been among the most intriguing of
prehistoric animals. N a m e d for the M e u s c River, that specimen gave its name
to all the mosasaurs that followed it into the limelight of paleontology. T h e
mosasaurs arose, diversified, and flowered 25 million years before the endCretaceous event and departed with the last of the terrestrial, nonavian
dinosaurs. M o s a s a u r fossils have been found in Canada, the Netherlands,
Sweden, Africa, Australia, N e w Zealand, Antarctica, and, in the U n i t e d
States, in Kansas, Nebraska, N e w M e x i c o , Colorado, Texas, Georgia, A l a bama, N o r t h and S o u t h Dakota, M o n t a n a , Arkansas, and N e w Jersey. T h e
rocks have revealed something of the lives of the mosasaurs, but much of their
existence remains hidden. T h e y may have descended from terrestrial lizards;
they may have given rise to snakes; but during their brief reign, they rose to a
position of marine dominance that would not be equaled until the whales and
dolphins arrived on the scene 50 million years later. S o m e reached e n o r m o u s
sizes, and with their flexible jaws and powerful teeth, they became the apex
predators of the open seas. Others lurked in the shallows, ready to ambush
anything that happened by. Still others developed heavy, rounded teeth that
enabled them to crush the thick shells of bivalves. T h e y may or may not have
punched through ammonite shells with their teeth. T h e y were big, fast,
powerful, and dangerous to other M e s o z o i c marine life-forms. Entombed in
the rocks, they left us tantalizing glimpses of an ancient way of life in the
water.
credence to the impact theory, because up to that time, the only evidence to
support it had been the enigmatic iridium particles and the disappearance of
so many species. T h e C h i c x u l u b Crater ( n a m e d for an ancient M a y a n village
near the site and pronounced C H E E K - z h u - l o o b ) , which is buried under
about 600 feet of sedimentary rock that has accumulated since the impact, is
125 to 185 miles across and was known to M e x i c a n petroleum engineers long
before anyone connected it with the impact. S u p p o r t e r s of the Alvarez et al.
(1980) theory of dinosaur destruction and there are many believe that the
impact induced a global environmental collapse of such magnitude that it
culminated in biological devastation. T h e y believe that the uppermost layers
of rock where the asteroid hit were heated to such temperatures that carbon
dioxide and sulfate aerosols were released into the atmosphere, creating a
worldwide climate of acid rain and smog, not to mention the darkening
effect of dust clouds circling the earth, which would lower temperatures drasticallythe so-called impact winter. Fiery debris ignited continent-sized fires
that burned for years. A cataclysm of this m a g n i t u d e brings out the hyperbole in writers, even if they are scientists. In Night Comes to the Cretaceous, James
Powell wrote: "A few minutes later, the mixture of vaporized meteorite and
rock, still traveling at ballistic velocities of 5 k m / s e c to 10 k m / s e c , began to
reenter the atmosphere. T h e individual globules were traveling so fast that
they ignited, producing a literal rain of fire. Over the entire globe, successively
later the greater the distance from the target, the lower atmosphere burst
into a wall of flame, igniting everything below. . . . Everything that could
burn did."
It is hard to resist such persuasively purple prose, but there are those who
have managed to do so. T h e y say that meteorites approach the earth all the
time, and although some have actually landed with a significant impact, they
have had little effect on the earth's atmosphere. And although the 1815 eruption of the Indonesian volcano Tambora may have darkened the skies and
killed off the year's corn crop in N o r t h America, it otherwise did no permanent damage. Besides, they say, the fossil record does not show that the land
dinosaurs died within a couple of years; rather, they died over a much longer
period, perhaps tens or even hundreds of thousands of years. T h e r e is little
doubt that a meteorite struck the earth about 65 m i l l i o n years ago, because
Triceratops,
and the great Tyrannosaurus were the last remnants of the line of giant reptiles
that had dominated the land for 150 m i l l i o n years. W h e n the asteroid hit, it
was these dinosaurs that were somehow eliminated. After the impact, no
terrestrial dinosaurs have been recorded. For the most part, their fossils have
been found in the H e l l Creek region of M o n t a n a . A l t h o u g h this does not
mean that M o n t a n a was the last refuge of the last of the dinosaurs, it does
mean that we have not found m a n y fossils elsewhere. It was also at this time
that the last of the mosasaurs died out. By the time the asteroid struck, the
ichthyosaurs and the plesiosaurs had been extinct for 20 m i l l i o n years.
T h e demise of the terrestrial dinosaurs is probably paleontology's greatest
mystery. For every detective who believes he has found the culprit, there is another who has identified a different suspect or g r o u p of suspects. But despite
the efforts of the best minds, the earth has kept the answerif there is an
answer hidden in the rocks. A concise s u m m a r y can be found in Michael N o vacek's 1996 book Dinosaurs of the Flaming Cliffs, in which the author, one of the
American M u s e u m ' s "dinosaur hunters" of the Gobi Desert in the 1990s, wrote:
Does that mean the long-standing mystery what killed the dinosaurs?
has been solved? W e l l , not entirely. Even if such an impact d i d occur at
sixty-five m i l l i o n years before present (give or take a h a l f m i l l i o n ) , we
cannot be sure it had the global impact ascribed to it. T h e r e is some
evidence that a drastic decline in n o n - a v i a n dinosaurs may have occurred
well before the end of the Cretaceous. Moreover, this devastation was
neither so overwhelming nor so rampant as the extermination at the end of
the Permian. True, m a n y families of m a m m a l s , birds, fishes, ammonites,
belemnites, and bivalves (clams, oysters, and other dual-shelled species)
went extinct. Over fifty percent of various marine planktonic groups were
also erased. But numerous important lineages many of the frogs, salamanders, turtles, lizards, m a m m a l s , crocodiles, birds, fishes, angiosperms,
conifers, arthropods (insects, spiders, crabs, and s u n d r y ) , gastropods
(snails and k i n ) , echinoderms, and nearly half the plankton species went
right on through the Cretaceous boundary. One of the real mysteries of
Cretaceous extinction is therefore its taxonomic selectivity.
W h y were the n o n - a v i a n dinosaurs so persecuted? A n d if such an event
was caused, as currently argued by many scientists, by a cataclysmic impact
of a giant asteroid, why was this pattern of extinction so discriminating?
W h y d i d some dinosaurs, feathery, flying creatures of high metabolism
that surely w o u l d not do well in clouds of m e t a l l i c vapors, survive the
event? W h y did many other vertebrates, animals, plants, and marine organisms m a k e it? T h e fact that this extinction event, like the Permo-Triassic
extinction and other events before it, was selective really complicates the
theory. It forces us to consider the subtleties of cause and effect relating to
the survival of biological systems. Namely, we need to know how exactly
such an event selectively snuffed out Cretaceous species. T h e s e are subtle
connections for which we have few insights.
Unlike the cataclysmic demise of the dinosaurs, there was no single event
to mark the departure date of the great marine reptiles. Also, when the
Niobrara and the Tethys Seas were closed by tectonic forces and dried up,
their water-dependent inhabitants dried up too. It wasn't like pulling the plug
from a bathtub, of course; the d r y i n g up and reduction of the inland seas took
millions of years, and during that time, many marine reptiles became extinct
without succumbing to evaporation. Moreover, not all the reptiles lived in
bodies of water that disappeared; many lived in oceans and seas that were
more or less where they are today. T h e marine reptiles passed into extinction
over a 20 millionyear stretch, the ichthyosaurs departing slowly, and the
plesiosaurs and the mosasaurs m a k i n g their final curtain call around the K - T
boundary. T h e C h i c x u l u b impact certainly wreaked ecological havoc on the
oceans, raining deadly chemicals on the waters, disrupting food chains, generating massive tsunamis, and affecting virtually every form of marine life.
T h e latest known plesiosaur fossil has been dated as late Cretaceous
( m i d d l e Campanian to M a a s t r i c h t i a n ) , about 65 million years ago: on S e y mour Island, Antarctic peninsula, fragmentary remains of an unidentified
plesiosaur were found (Chatterjee and Zinsmeister 1982), along with the partial skull of a very large mosasaur, provisionally identified as either Hainosaurus
or Tylosaurus. M a n y elasmosaur fossils are known from the late M a a s t r i c h t i a n
rocks of the Maastricht region of Belgium. M o r e recently, remains of elasm o s a u n d plesiosaurs have been collected from the lower part of the late
Cretaceous L o p e z de Bertodano Formation on S e y m o u r Island, Antarctica,
by the Polish Antarctic Expeditions (Fostowicz-Frelik and Gazdzicki 2001):
the not fully grown elasmosaur, which m a y have lived in shallow water
environment. T h e studied remains share some similarities with those of
Mauisaurus from the M a a s t r i c h t i a n of N e w Zealand.
the largest white shark teeth are a little more than 2 inches long, fossil
Megalodon teeth can be 6 inches. Megalodon (which means "great t o o t h " ) is
extinct, gone from the oceans for at least 100,000 years, but it is difficult to say
why. It was obviously a powerful predator, and it is known that it preyed on
whales, but the whales are still here and the predator is long gone. T h e watery
climate may have changed in such a way that w a r m - b l o o d e d m a m m a l s could
survive but cold-blooded reptiles and fishes could not; or perhaps the prey
species of the great marine reptiles gradually disappeared, which would have
sent the predators drifting hungrily toward extinction. But because we really
don't know what the primary prey of the large marine reptiles was some
theorists hold that it was smaller marine reptiles the question remains unanswered, and maybe unanswerable.
T h e only extinctions we truly understand are those that we ourselves
engineered: the great auk, dodo, passenger pigeon, Carolina parakeet, quagga,
Tasmanian wolf, Caribbean monk seal, Steller's sea cow, and hundreds of
others, all eliminated in the brief moment that we have had to strut and fret
our apocalyptic hour upon the evolutionary stage. As for why so many other
species disappeared long before M r . Sapiens arrived on the scene with his
clever weapons and his special kind of havoc, let us give the final word on the
subject to Charles Darwin:
We need not marvel at extinction; if we must marvel, let it be at our own
presumption in imagining for a moment that we understand the many
complex contingencies on which the existence of each species depends. If
we forget for an instant that each species tends to increase inordinately, and
that some check is always in action, yet seldom perceived by us, the whole
economy of nature will be utterly obscured. W h e n e v e r we can precisely say
why this species is more abundant in individuals than that; why this species
and not another can be naturalised in a given country; then, and not until
then, may we justly feel surprise why we cannot account for the extinction
of any species or g r o u p of species.
References
Adams, D. A. 1 9 9 7 . Trinacromerum bonneri, new species,
9(') 47-'5:
358:507-14.
London
126:40347.
Press.
80:62742.
Museum
(Natural History),
Academic Press.
Archibald, J. D, and W. A. Clemens. 1 9 8 2 . Late
London,
Part
I.
British Museum.
. 1 9 1 2 . Description of a new plesiosaur
Accademia
malematiche e
nazionale
dei
naturali
Lined.
Classe
di
scienze
fsche
5zf j):5j56.
London 7 8 : 2 8 5 9 8 .
13(2)121-34.
mammals. Evolution 2 5 : 6 3 6 5 8 .
232(4)58-78.
1986.
3(2)1122.
Natuurhistorisch
Cenootschap
in
Limhurg
41(1)5255.
85.
Doubleday.
Bardet, N. 1 9 9 2 . Stratigraphic evidence for the
extinction of the ichthyosaurs. Terra Nova 4 : 6 4 9
56.
80:1114.
Historical Biology 7 : 3 1 3 2 4 .
vol.
2.
Technical Report N P S / N R P O / N R T R - 9 5 / 1 6 ,
pp.
Plenum.
56:52024.
Paleo.
34-39.
Scientist
i}2(i79o):4o44.
Press.
. 1 9 9 7 b . Vertebrate Paleontology. Chapman and Hall.
43 534-3 -
western
Kansas.
Kansas
Ceological Survey,
Current
Bulletin 2 4 4 ( 1 ) : ! 2 6 .
0325:36986.
74-
7:332-33-
37(4^:94153.
Palaeontology
langhami Brown
Paleo. 1 7 : 2 9 5 3 0 7 .
uesd.edu.paleontology/seaway.html.
Curious
Things
of Nature
Existing
before
Linn. Soc. 1 2 5 : 1 1 5 4 7 .
14:5765.
Biology
Historical
Casinos.
of the Prehistoric
World.
Fourth Estate.
Caldwell, M. W. 1 9 9 4 . Developmental constraints
'4(4>459-7'-
Dinilysia.
Acta Palaeontol
Polon. 4 6 ( 2 ) 2 0 3 1 8 .
Vert. Paleo.
22(4):86i66.
178:3758.
1989b.
Mexico.your: Paleo. 6 3 ( 6 ) : 9 3 0 3 9 .
Kolposaurus Camp,
35(6)822.
Association of Canada.
Callaway, J. M . 1 9 9 7 a . Ichthyosauria. In J. M .
preoccupied. Jour.
Paleo.
228(2): 3644.
K.
1989.
Dolichorhynchops
Trinacromerum.
201(2)125987.
2(2):i4873.
constraint.
Paleobiology
13:32641.
Zoologist
17(5>66.
31:64454.
London 2 9 7 : 3 1 5 8 3 .
1988.
Life Books.
Colbert, E. H. 1 9 4 9 . A new Cretaceous plesiosaur
London B 5 2 5 : ^ 8 7 4 0 0 .
18:299318.
I4(i):i42.
69.
15(12)503-7.
Paleo.
22(3)629-44.
12(1)66-86.
Geology
Modern
Novitates
.
2076:116.
1963.
Dinosaurs:
I heir
lie//,/.
Hutchinson.
. 1 9 6 5 . The Age of Reptiles. W. W. Norton.
. 1973.
. 1 8 7 0 b . On
Amer. Jour.
Sci. 5 o ( i 4 8 ) : i 4 o 4 1 .
Soc.
11:27174.
I 'ress.
Conway Morris, S. 2 0 0 2 . W e were meant to be. . . .
New
Scientist
lyb^z^bgy.zb29.
(l8jl)
Territories,
Report
Trans.
Ceol. Soc.
pp.
London i ( 2 ) : i o 3 2 3 .
2:38190.
4 0
. 1 8 7 2 b . [On the structure of the Pythonom o r p h a ] . Proc. Acad. Nat. Sci. Phila. 2 4 : 1 4 0 4 1 .
57)^63-64.
Soc. 1 2 : 2 6 4 8 7 .
Nat. 3:8491.
66.
1869c.
Ormtholarsus
Remarks on
nnmams,
and
Paleobiology, pp.
Thoracosaurus brevispinus,
Macrosaurus
proriger.
Proc
Press.
. 2 0 0 0 . History of Lije. Blackwcll Science.
1. http: / / www.dinosauria
Dino-Dispatches No.
.com/dispatches/ 19981108001.html.
64.
Chicago Press.
. c o m / d m l / n a m e s / mosa.html.
Pronunciation Guide, h t t p : / / w w w . d i n o s a u r i a
Lower Jurassic
megacephalus,
Knight.
Dutton.
Darby, D. G., and R. W. Ojakangas. 1 9 8 0 . Gastroliths
from an Upper Cretaceous plesiosaur. Jour. Paleo.
54:548-56.
and Brothers.
DeBraga, M., and R. L. Carroll. 1 9 9 3 . T h e origin of
mosasaurs as a model of macro-evolutionary
patterns and processes. W. Biol. 2 7 : 2 4 5 3 2 2 .
Palaeontology
45(?) 55775:
18:42342.
5=559-94-
1982.
18:6181.
21:50439; 22:70112.
5:99107.
381:1-157.
. 1906.
23:983.
Eldredge, N.
1991a.
Evolution. Prentice-Hall.
. 1 9 9 8 . The Pattern of Evolution. Freeman.
. 2000.
Creationism. Freeman.
Ellis, R. 1 9 9 9 . The Search for the Giant Squid. Penguin.
Erickson, G. M., K. C. Rogers, and S. A. Yerby. 2 0 0 1 .
growth rates. N a t u r e 4 1 2 : 4 2 9 3 3 .
3):45A-46A.
N.S.W2:41013.
. 1 8 9 7 . An Australian sauropterygian
Aust. Mus.
Geosei. 8 i ( i ) : i - 8 .
Douglas, K. 1 9 9 9 . Dinodolphin. New Scientist
i6i(2i76):i4.
3:2129.
5:30616.
Muraetwsaurus
Seeley.
Mercian
Geologist
i4(4):i9i- 6.
Vert. Paleo.
i7(suppl.
to j):4jA-44A.
Prehistoric
Times 4 4 : 2 9 3 1 .
I4(abstracts):i9.
Fastovsky, D. E., and D. B. Wcishampcl. 1 9 9 6 . The
[.volution and Extinction of the Dinosaurs. Cambridge
University Press.
Fernandez, M. S. 1 9 9 4 . A new long-snouted
104(1-2)56-75.
7:497-84.
36.
): 2A.
Morphol. 2 2 5 : 5 1 6 0 .
270
5EA
D%A OTVJ
. 1998.
Ensign.
Gilmorc, C. W. 1 9 1 2 . A new mosasauroid reptile from
the Cretaceous of Alabama. Proe. U.S. Natl. Mus.
14(4)734-41.
4'( 7)479- 4l 8
Part 111.
Polon. 6 0 : 7 3 2 .
Palaeontol.
164:19394.
Paleobiology 3:11551.
Sept. 192.2, p. 3 0 .
1988.
Ameghiniana 2 5 : 3 1 6 .
History.
Ameghiniana 3 0 ( 3 ) 2 4 5 5 4 .
Worldwide.
BBC
Geol. Assoc. 1 0 7 : 1 0 7 1 6 .
3 5 437-552
Freeman.
. 1 9 7 1 . Liopleurodon rossicus (Novozhilov) a
Palaeontology
14:56670.
London 1 4 6 : 3 7 4 0 .
Holmes,
R.
1996.
Plioplatecarpus primaevus
Sterling.
Harlan, R. 1 8 3 4 a . Notice of the discovery of the
Batrachiosaurus.
Hawkins,
T.
Proc.
Ceol. Soc.
London 3 : 2 3 2 4 .
Fletcher.
36:363-69.
London 1 0 6 : 3 1 8 2 1 .
Rev. 3 1 ( 3 ) : 1 4 8 - 5 1 .
resemble
Ichthyosauria).
those
of
Ornithorhynchus paradoxus.
Phil.
Western
Nostrand.
Evolution of the
National
Museum of Wales.
Biol. 2 0 5 : 4 3 9 4 1 .
Hill.
26(10)947-50.
London B 3 5 2 : 5 6 9 1 .
Vert.
2i(suppl. to 3 ) 6 7 A .
Linn. Soc. 6 5 : 3 6 9 4 5 3 .
Geology
18:44358.
74(1)915-37.
Palaontol. Z. 6 8 ( 1 2 ) 2 5 9 6 5 .
Nature
400:65559.
intermedins,
Lciodon
proriger,
Proc.
Baptemys
Acad.
wyomingensis,
Nat.
Sci.
Phila.
22:3-5.
4o(i):916.
PalAsiatiea 4 o ( 2 ) : i i 4 2 6 .
Geology
16:22948.
Palaeontology ^ 4 . ( ^ ^ : 6 5 ^ 7 0 .
i8(4):70917.
31:30810.
. 1 9 9 4 a . 1 he durophagous mosasaurs
(Lcpidosauromorpha, Squamata) Globidcns and
Carinodens from the Upper Cretaceous of Belgium
and the Netherlands. Paleo. Jour. 3 3 ( 6 ) : 6 ^ 8 4 6 .
Mosasauridac)
Palaeont. Abh.
?):^5574.
62:17194.
London B 2 6 6 : 2 3 6 7 7 3 .
34:287502.
. 2 0 0 2 a . Extinction of ichthyosaurs: A
1930.
1932.
Mem.
Qld.
Restoration
of Kronosaurus tjucenslandicus.
Mus. 1 0 : 9 8 .
3(6)388-90.
1998(7)401-14.
Nat. Belg. 5 9 : 1 3 7 9 0 .
Palaeont. Abh. 2 1 7 ( 1 ) 1 2 5 .
Press.
Long, |. A., and A. R. I. Cruickshank. 1 9 9 8 . Further
1972
from
Aust. Mus. 1 9 : 4 7 5 5 .
1998(1)2641.
Abb.
209:10534.
I.ethaia 3 3 : 7 1 7 4 .
44(6)112756.
Kaupia
2:7797.
Palacontological
Association.
220(3)431-47.
38:897903.
the Evidence:
How Did
They Know
That?
BBC.
Martin, J. E. 1 9 9 4 . A baby plcsiosaur from the late
Germany.
Palaeontology
43(1)2940.
77:4.667.
on File.
13352.
Paleo.
. 1 9 7 2 b . T h e systematics of Cretaceous
101(9)48-53.
19:355-56.
7(2):i2i37.
prcdation.
1 9 9 4 . Cymbospondylus (Ichthyosauria:
11(1)9-29.
communis,
I.
breviceps,
and
Stenopterygius
Roy.
Ontario
Mus.
93:121.
Mus.
100:130.
97:i-37-
Sciences, ser. 2, 3 1 3 : 1 2 0 7 1 2 .
3(5)668-83.
5(3)28595.
1,2.2.
6078
45456.
Vert. Paleo.
9(3)269-81.
ichthyosaur Eurhinosaurus
Palaeontology
. K)94d.
longirostris.
37(4)747-53.
Tcmnodontosaurus riser is a juvenile form ot
T. platydon. Jour.
Vert. Paleo. 1 4 : 4 7 2 7 9 .
. 1 9 8 9 b . T h e lehthvosaunan tailbend: A
1989c.
35 555-7o-
tomography.
Paleobiology
Paleo. 5 2 : 1 1 5 5 6 2 .
Paleobiology
15(4)142936.
Lias.
32:292-303.
Palaeontology
32(3)40927.
Earth Sci. 3 3 : 4 3 9 4 3 .
University Press.
. 1 9 9 1 b . An ichthyosaur forcfin from the Triassic
pp.
Reptiles,
2001.
The
Grippia
Academic Press.
6180.
Dragon
Ancient
Seekers.
ichthyopterygian forefins.
Vert.
Zoo/.
Jour.
London
1998.
Ceo!
Univ.
sea.
Oceans
Jour.
Vert.
Paleo.
Skull of
2000c.
283(6)5259.
American
Grippia
No
longirostris:
ichthyopterygia.
Palaeontology
43(1)114.
"Nature 4 1 5 : 3 0 9 1 2 .
chaoxianensis
(Ichthyosauria)
shows
Jour.
Paleo.
35(897)377-78.
Science
Utatsusaurus
72(0:133-36.
. 1 9 9 8 b . Taxonomy and limb ontogeny of
hataii.
Chaohusaurusgeishanensis
Vert.
Paleo.
hataii
(Ichthyosauria)
Jour.
0 )
status of
Utatsusaurus
Lethaia
30:22128.
Jour.
Vert.
Paleo.
17(1)3944.
Himalayasaurus
tibctensis
Ichthyopterygia
2(2)17481.
Grippia
Paleo.
5 - 4 -
Paludicola
longirostris
Vert.
ichthyosaurian specimens.
Jour.
Paleo.
7<3>475-79-
1997c.
with
Utatsusaurus
(Ichthyosauria),
16(3)396402.
Scientific
11(2)5356.
Jour.
ichthyopterygian? A
5:38190.
Calif.
Omphalosaurus
Is
2000a.
Bull
four.
19(3)47295.
ichthyosaurs.
Wiley.
Revolution.
19(1)28
Paleo.
20(2)295
199:129.
The Evolution
Vert.
Paleo.
phylogenetic perspective.
19(1)4249.
Paleo.
1999b.
Vert.
four.
41.
42(5)92752.
Shonisaurus.
41(4)59199.
Perseus.
Palaeontology
longirostris
Palaeontology
Ancient Marine
Nature
393:25557.
Nature
402:747.
Nature
382:347-48.
Can.
Jour.
Earth
Sci.
38:983
tool.
27(4)72434.
Paleobiology
Triassic-Jurassic gap.
evidence.
Bull.
Inst.
Roy.
Sci.
Nat.
Dinosaur
World
(Reptilia:
Phalarodon
mixosaurs.
Palaeontographica
o f
Li,
Qianichthyosaurus
1999
(Reptilia,
3:10324.
Triassic ichthyosaurs.
from
of
252:122.
material
Evolution
interrelationships
4:98103.
Press
reptile
New material of
4:27-29.
Ditto
Palaeoecology
79:149-69.
1999.
1998.
Palaeocltmatology,
Belg.
70:16178.
Naish, D.
Palaeogcographv,
Animals
io(2):6i63.
Jour.
Vert.
Paleo.
22(4)175965.
Can. Jour.
Hainosaurus
(Reptilia:
Can.
Jour.
Laccrtilia)
Earth
Sci.
from
of
Utatsusaurus
(Reptilia:
Sci.
University
o f
Copenhagen
1994.
Novacck, M . J.
Prehistoric
1996.
Life.
Macmillan.
Dinosaurs of
the
Flaming
Cliffs.
Anchor Doubleday.
Obradovich, J. D. 1 9 9 3 . A Cretaceous time scale. In
30:48690.
of Derby.
Norman, D.
3840.
Jour.
pp.
Life
North
25:156470.
Water,
to
America.
in
Adaptation
Gcologisk Museum.
13:185-88.
Sci.
Secondary
Vert.
Paleo.
14:450-52.
of
the
Western
Interior
pp.
Basin,
37996.
Evol.
Biol.
I4(6):g8791.
Acta
Zool.
Fcnnica
213:1
Soc.
28(i):ioi12.
Paleobiology
Jour.
Vert.
3> 3A.
Natural
History
no(5):22-24.
Amer.
Mus.
Nat.
Hist.
Mem.
Sphenodon.
Osborne, R.
Amer.
Nat.
1998.
The
Ostrom, J. H.
Pliosaurus
Monog.
111:112.
London
Floating
Egg:
in
the
Making
1969.
Osteology of
of
Soc.
Bull.
Plesiosaurus
Peabody
Mus.
18401845.
London
macrocephalus
(Conybeare).
for
the
Advancement
London
33:133
of the
Prehistoric
World.
Discovery
E.
2000.
Yaguarasaurus
columbianus
Historical
Biology
14:12131.
Evolution
of the
Western
Geological Association
31932.
Twenty-ninth
of
Science.
Nature
398:50813.
Ichthyosaurus
New
Scientist
i73(233o):g.
Proc.
Geol.
Meeting
John
of the
British
Murray.
Ichthyosaurus
(communis?).
Ann.
Mag.
Nat.
Hist.
17:44
46.
Perkins, S.
the
Atlas
M.
embryo of an
Hippolytc Bailliere.
3:2428.
of
Soc.
Report
1999.
uncovered.
i860.
Geol.
Carboniferous of Scotland.
5:51114.
Odontography.
Quart. Jour.
of Canada.
33:682715.
Nat.
2:66366.
London
London
Books.
Interior Basin,
London
Soc.
Geol.
Geol.
Deinonycbus
30:1165.
Geol.
Jour.
Episodes
specimen
Quart.
Colombia.
34(397)117.
Cretaceous of Montana.
Association
portlandicus.
Cape.
Ceology.
Soc.
Soc.
P.
and
trochanteric
Paramo-Fonseca,
Trans.
P.
Palmer, D.
i(4>i67-88.
Proc.
38.
7(i82):84i45.
and cartilaginous.
Hist.
Palaeont.
2728.
Sauropterygia.
antirrbopus,
Monog.
of
grandis.
Science
Pliosaurus
pp.
London,
Palaeont.
(suppl. to
22
Paleo.
Clay,
63.
2002.
Sea dragons.
Science News
i62(8):i22
24.
Sauropterygia). M e m . Q l d . M u s . 2 0 : 6 4 7 5 5 .
Secondary
Adaptation
Powell.
Mammals
pacificus?
Mar.
Mam.
1994.
Coniasaurus
crassiiens
Jour.
\erl.
suppl. to
14
Paleo.
Powell, J. L.
1998.
11:37688.
Night
Comes
to
the
2002.
of
the
Cretaceous.
Freeman.
pp.
119.
Trans.
Roy.
Soc.
London
1991b.
1994.
pp.
of Dinosaurs,
56273.
Biol.
1994.
Osteology
of
Simosaurusgaillardoli
and
the
28:185.
Ancient
Marine
Reptiles,
pp.
107
. 1 9 9 8 . Ichthvosaur r e m a i n s Reptilia.
325:42135.
Extinction:
Bad
Genes
or
Bad
Norton.
Luck?
Acad.
Sci.
Proc.
Mh.
N.
]b.
Geol.
1998:53744.
2001a.
Jaekel,
1902
Geol.
Science
Placochelys
Cyamodontoidca (Reptilia,
91:6758-63.
Placodonta).
Fieldiana
45:1104.
2001b.
A new species of
Nothosaurus
(Reptilia:
215:1501-3.
231:83336.
Paleontology.
placodonta
Science
Marine
I3i(i786):4649.
Scientist
Natl.
to
22:37130.
Evol.
Palacont.
Guide
Society
1 9 . Academic Press.
Encyclopedia
E. L. Nicholls, eds.,
University of
(Chicago Press.
extinction.
Audubon
Knopf.
National
World.
Padian, eds.,
Fieldiana
173:17.
Nitecki, cd.,
University
70.
Academic Press.
p.
Water,
K.
:42.x.
Dorset.
in
15(^^:5 ji49.
Sci.
Life
to
1971.
Principles of
Freeman.
263:13761.
Science
219:124041.
Ancient
Marine
Reptiles,
pp.
12144.
Academic Press.
Ricppcl, O., and R. R. Rcisz. 1 9 9 9 . The aquatic
origin of turtles. In E. Hoch and A. K. Brantsen,
eds.,
Secondary
Adaptation
to
Life
in
Water,
p.
59.
mosasaurs.
Riess, j . 1 9 8 6 . Fortbcwcgunswcisc.
236:7577.
Science
ichthvosaurer.
Paleontographica
192:93155.
Kansas
Ceol.
164:19394.
Palaeont.
Riggs, E. S.
1944.
Univ.
Rudwick, M . ] . S.
Aust.
Aust.
23(7)538-45.
Hist.
Jb.
Ceol.
N.
Jb.
Ceol.
Palaeont.
Amer. four.
Osteology
of the
Reptiles.
University of
Chicago Press.
.
1966.
Paleontology.
University of Chicago
1968.
The
Procession
of Life.
World.
or secondary? A n n . S. Afr. M u s . 6 4 : 2 2 1 3 0 .
Kronosaurus.
from
Deep
Time.
University o f
Ceorgcs
Cuvier,
Fossil
Bones,
and
Geological
Postilla
Yale
Univ.
Bull.
Peabody
Mus.
Nat.
Hist.
zy.iz$j.
Ceol.
1975a.
3(7)36980.
Mem.
A new species of
from South
Clobidens
American
33(13)23556.
Ceol.
of North
The
America,
Cretaceous
pp.
System
11936.
in
the
Western
Geological
Association of Canada.
112:115.
1997.
Fieldiana
Breviora
of Fossils.
Press.
.
Fieldiana
Vertebrate
Meaning
246(2)10921.
1956.
Scenes
American mosasaurs.
153:86128.
Iehthyosaur ancestors.
1948.
1992.
mosasaurs.
149(3)286332.
Palaeont.
Catastrophes.
The
Chicago Press.
1990.
1972.
19:40813.
Hist.
22(8)368-70.
Hist.
Nat.
Nat.
1988.
Nat.
Aust.
Palaeontologia
http://palaco-electronica.org/
3(1).
Electronica
2 o o o _ i / r e t i n a l / other.htm.
30:7787.
Bull.
19(2)7382.
Canada
N. Jb.
Ceoscience
246(4)96105.
1989.
Crocodiles and
1989.
America.
Alligators.
Facts on
File.
Rothschild, B. M., and L. D. Martin. 1 9 8 7 . Avascular
An
Odyssey
in
Time:
The
Dinosaurs
of
North
Kaufmann,
eds.,
Evolution
of
the
Western
Interior
Basin,
1989a.
buseheri,
Science 2 3 9 : 7 8 0 8 3 .
T h e large ichthyosaur
Vert.
ate ammonites.
9(2)a6373.
Paleo.
. 1 9 8 9 b . T h e pachypleurosaurids (Reptilia:
Trans.
Roy.
Soc.
London
1992.
Pleistocene serpent
Wonambi
Nature 4 0 3 : 4 1 6 2 0 .
325:561670.
.
Paleo.
394:62930.
Nature
evolution of snakes.
of a new species.
four.
Cymbospondylus
Pachyraehis.
Llistorical
13:12752.
Biology
Cymbospondylus
1997.
Sijualieorax
Palaios
12:71-83.
Jour.
Paleo.
1997.
66(2)33237.
T h e paleobiogcography of
Reptiles,
pp.
1743.
In
Shastasaurus.
Ancient
Z.
Seeley, H. G.
(Ichthyopterygia:
Soc.
.
Reptilia) in the
Jour.
Paleo.
64(i):i6i
N. Jb.
Geol.
Palaeont.
On
London
On
1877a.
Q. Jour.
Q. Jour.
Geol.
Soc.
33:54146.
London
bones of
Pliosaurus
evansi
Jour.
Geol.
Soc.
London
30:716
3-
Jour.
Ichthyosaurus
Br.
Assoc.
Adv.
50:68-76.
young in
controversies.
1887:697-98.
18:559.
Geol.
Seeley, an
Mauisaurus gardneri,
certain species of
Geology
leedsi,
Modern
Muraenosaurus
30:197208.
10:63140.
Paleo.
Can.
Vert.
Bd.
Mh.
1874.
Folkestone.
64.
Res.
Lisb.
74(1
2)1-35.
of
in the
Xiphiasgladias,
four.
25(5>9?-'9-
Academic Press.
Plesiosaurus.
Rep.
Br.
Assoc.
Adv.
Sci.
Sci.
Ann.
Mag.
Nat.
8(r):4j641.
Hist.
Ann.
Mag.
Nat.
8(1)43641.
Hist.
Placenticeras
Geowissenschaftliehe
Trans.
Kansas
. 1879. T h e scales of
1:93102.
Jour.
Vert.
Sci.
6:54
58.
Acad.
Liodon.
Amer.
Nat.
Goronyosaurus
13(2)132.
from the
Paleo.
31:74762.
Palaeontology
7(suppl. to 3):25A.
. 1 9 8 9 . Implications of juvenile mosasaur
recognition on taxonomy.
Jour.
Vert.
Paleo.
(P.
9(suppl.
and
to 3 > 3 8 A .
Jour.
Vert.
Paleo.
io(suppl. to
conyheari)
with observations on
P.
hrachycephalus,
megaloeephalus,
Stutchberry,
four.
Soc.
P.
Owen.
Geo!
London
37:472-81.
Spotila, J. R. 1995. Metabolism, physiology, and
thermoregulation. In K. A. Bjorndal, ed.,
3> 2A.
4
Vert.
Paleo.
I2(suppl. to
3):5iA.
Ancient
Marine
Reptiles,
pp.
Conservation
of Sea
pp.
Turtles,
Biology
59192.
and
Musick, eds..
33354.
The Biology
of Sea
Turtles,
pp. 297314.
C R C Press.
Academic Press.
ed.,
Paludicola
Press.
2(2)190205.
mosasaur,
Plioplatecarpus
primaevus.
Jour.
Vert.
Paleo.
Press.
Bull.
Geo!
58:1263.
Simpson, G. G. 1953.
. 1987.
Trans.
Kansas
Acad.
Sci.
20:12224.
1907.
Yale University
Trans.
. 1909.
.
Extinction.
2:133.
Soc.
Scientific
250(6)6472.
American
i6(suppl. to j):64A.
Methodist
Extinctions,
Deer
Kansas
The
1017.
Hunting
River,
Alherta,
Acad.
Life
of a
Sci.
Fossil
Dinosaurs
Canada.
21:111 14.
Hunter.
m
the
Henry Holt.
Hid
Lawrence,
Lands oj
the
Red
Kans.: n.p.
Trans.
Kansas
Acad.
Sci.
30(1)11920.
Somerset, England.
Vert.
Paleo.
9:j9A (abstract).
Sci.
Nat.
Hist.
Mus.
L A .
1984.
Elasmosaurus
and a page
platyurus
17(2)2527.
Discovery
Peabody
Corosaurus
Nat.
Hist.
Amer. Jour.
Paleobiol.
7:217
PPlesiosaurus
sp.
Arcbaeol.
Soc.
England.
Proc.
Dorset
Nat.
U.K.
Quarterly
116:7276.
Dinosaur
1(2)67.
Plesiosaurus.
In J. M .
Ancient Marine
. 1 9 9 9 . An examination of plesiosauria
(Diapsida: Sauropterygia) from the Niobrara
Chalk (Upper Cretaceous) of central North
America.
286
S A
Univ.
D'RA
Kansas
Paleo.
OTv\C
Trans.
Roy.
30:69368.
London
Placodusgtgas
Jour.
Vert.
and
Paleo.
7:138-44.
Pistosaurus
and
Zool.
Jour.
Linn.
Soc.
London
90:10931.
Crocodiles
and
Alligators,
Nature
346:14
[5.
. 2001. Ruffling feathers. Atewrf 410:103637.
Natural
2:812.
History
and
Hist.
Russia
67(12)1779 -
Dorset,
in
Silvestrosaurus
Charmouth,
of Dinosaurs
28.
Z.
Age
Palaontol.
The
293-A:6390.
Rev.
Mongolia,
Soc.
44:1151.
16:40320.
Paleo.
Bull.
Sci.
Vert.
Kurichkin, eds.,
Storrs, G. W.
Jour.
County
441:1-9.
aleovensis
teleosts.
Contrib.
11:115.
Nat.
Hist.
Mag.
2:27175.
Ceol.
Surv.
Nigeria
Bull.
1:4248.
Ann.
Mag.
Nat.
Hist.
10(6)2069.
. 1931.1 he plesiosaurs in the Bristol Museum.
Rep.
Br.
Assoc.
Adv.
Sci.
1930:34041.
Bristol
Nat.
1973.
Tarlo, L. B.
1958.
Phillips.
.
1959b.
and
Pliosaurus
Museum.
Pliosaurus
macromerus,
(Owen)
from
the
Palaeontology
2(1^:3955.
Br.
Mus.
(Nat.
Hist).
Geol.
N. Jb
Geol.
Palaeont.
Z.ool.Jour.
Linn.
Soc.
112:17996.
London
1995.
Natural
History
Reptiles,
Ancient
Marine
1876.
The Yorkshire
Lias. Van
strategics
by
aquatic tetrapods.
Historical
Biology
Nature
anatomy and
290:62829.
Lifestyle of plesiosaurs.
1986.
Nature 3 1 9 : 1 7 9 .
Zoo! Jour.
Soc. 9 1 : 1 7 1 9 5 .
swimming and
buoyancy.
30(33:55135.
Palaeontology
1989a.
Soc.
New Scientist
1989b.
1989c.
Nature 3 3 8 : 3 8 1 .
Nature 3 3 8 : 6 3 5
Natural
Rhomaleosaurus
zetlandicus
Phil.
Trans.
341:399418.
104(10)6671.
History
36(2)35760.
2000.
Science
287(5460)201012.
36.
Pliosaurus
the
L. J. Jacobs.
121(1655)65.
aquatic
London
England.
T h e other dinosaurs.
from
. 1 9 8 7 b . A reinterprctation of ichthyosaur
Plcsiosauria)
Linn.
functional morphology of
(Reptilia:
brachyspondylus
large
London
14:15-31.
ballasting.
Soc.
Scientist
Roy.
Abb.
Voorst.
Trans.
164:18892.
Phi!
341:163-75.
164:19394.
Soc.
104(10)69.
(5):i47-8 .
Palaeont.
Roy.
i(4):28391.
Palaeontology
Trans.
1:19399.
brachyspondylus
Clay.
Stretosaurus
British
Amphibians.
I'hil.
335:24780.
London
The scapula of
Palaeontology
1959a.
Kimmcridge
.
27:34360.
Soc.
Fossil Reptiles
Paleobiology
2:1938.
locomotion
in
sharks.
American
Zoologist
Palaeontology
Modern
Geology
18:489501.
.
Jour.
Jour.
Sci.
Paleo.
Texas
2i(i):6979.
1981.
Fossil
Vertebrates
(Xiphiasgladius
L.):
Its
Fish.
Res.
Bd.
Can.
Mitteilungen
Center
Geol.
of the
1986
Earth.
10:37076.
Palaont.
1984.
Platypterygius
an Australian
australis,
Cretaceous ichthyosaur.
l.ethaia
17:99113.
Platypterygius
den
Iithograpfuschen
Akademie
der
1997.
1998.
Wissenschaft,
1992.
The
On
Call
Time
Gelehrte
Methuselah's
of Distant
Machine:
Aust. four.
Sci.
6:167-78.
schicfcrn
aus dem
Polyptyehodon
Bull,
der
Freeman.
Trail.
Scientific
koniglische
6:66170.
Anzeigen
Copernicus.
Mammoths.
Explorations
in
Deep
Copernicus.
2000.
Rare Earth.
Copernicus.
Watson, D. M. S. 1 9 2 4 . T h e elasmosaurid shouldergirdle and fore-limb.
Proc.
Zool.
to
Whales
Soc.
2:885
London
917.
Modern
Geology
18:5039.
Torrens, H. S. 1 9 9 5 . M a r y Anning ( 1 7 9 9 1 8 4 7 ) f
Lyme: "The greatest fossilist the world ever knew."
Hist.
Miner.
Wade, M .
Tune.
Innsbruck
20:34969.
four.
Zbl.
Ward, P. D.
130:147.
Br.
the
Ichthyosaurus
to
28:115-37.
T h u r m o n d , J. T , and D. E. Jones.
of Alabama.
A fourney
longipinnate
42:128996.
1969.
1864.
25:351-59.
Texas.
Verne, J.
Sci.
Palaeoclimatology,
Palaeoecology
1981.
Sea
Guide
of the
World.
Hutchinson.
Weems, R. E., and R. B. Blodgett. 1 9 9 6 . The
pliosaurid
Megalneusaurus:
newly
recognized
28:25784.
ammonite
Watson, L.
Surv.
Bull.
169:24570.
Soc.
Geol.
Palaeogeography,
U.S.
2152:16975.
Amer.
Bull.
Geol.
50:1974.
mosasaur
and Colorado.
Platecarpus.
59(3)382-84.
Trans.
Kansas
Acad.
Sci.
Univ.
Calif.
Mem.
13:125254.
Shale of Texas.
Fondren
Sci.
See,
S.
Methodist
1:1
Univ.
28.
Univ.
Calif.
Pub.
Sci.
29:46-144.
Univ.
Calif.
Pub.
Sci.
Univ.
Wiffen, J.,
45:12.
Notes
1980.
Evolution 5 5 ( 1 1 ) 2 3 0 3 1 8 .
Moanasaurus,
7582.
Natural History.
Mus.
Dinosaurs of
Ceol.
46:196.
Nebraska
pp.
New Mexico,
. 1970.
1949.
Alzadasaurus
Jour.
Paleo. 2 3 : 5 2 1 3 5 .
Rec.
Canterbury
ga-
Mus.
Geol.
23:50728.
Geophys.
NZ. Jour.
Geol.
in.
33:6785.
Geophys.
Dinosaur
four.
Paleo.
573-33-
1959.
1968.
Before
Deluge.
Houghton Mifflin.
Doubleday.
(Clobicephala
Traill).
tnelas
Mar.
Mam.
Paleo.
1935.
On the skull of
queenslandicus
Longman.
Nat.
8:219-28.
1940.
Occasional
Boston
Soc.
Story
of New
Zealand's
Century.
N.Z. Jour.
Geol.
29:20552.
Geophys.
28:62540.
Geobios
1986.
The
Riddle
of the Dinosaurs.
Holotype of
Plesiosaurus
longirostris
Knopf.
Blake
Jour.
Paleo.
System
in
the
Western
Interior
of North
America,
The Cretaceous
pp.
14:451-67.
Random
Woman.
The
Kronosaurus
Papers
Ann.
Dragons:
Wilford, J. N.
51(2)1424-25.
the
Hist.
of
Sci.
16(2)299-325.
White, T. E.
Valley
Polyplychodon
Wendt, H.
1991.
Nat.
Hist.
9:74244.
the plesiosaurs.
Trans.
Kansas
Acad.
Sci.
13:12122.
Cretaceous of Kansas.
Trans.
Kansas
Acad.
Winchester, S.
Sci.
12:17478.
.
William
1890.
Science
Kansas mosasaurs.
1891.
Univ.
Kansas
Amer.
Nat.
Univ.
Quart.
Univ.
Geol.
Surv.
Restoration
Dolichorhynchops
Columbian
Kansas
Univ.
Sci.
Bull.
1906.
1907.
Polycotylus.
Amer.
T h e skull of
Jour.
Brachauchenius,
Sci.
Proc.
U.S.
Science
with special
1914.
Jour.
Nat.
52:47789.
Mus.
Reptiles
1992.
Ciidastes
liodontus
Vert.
8(5)54545.
Paleo.
1988.
Selmasaurus
8:1027.
Paleo.
loggers, your
Exp.
202(22)512126.
Biol.
the
The
Past
Diversity
Hist.
Soc.
1819.
and
Vert.
18:113.
Paleo.
of
Pachyhachis
and
problematicus,
Sci.
Paris
the
Comptes
Rendus
529:85157.
Present.
a
of Life.
Wernerian
5:45057.
Mem.
16:71555.
of
Harvard
mount
for
plesiosaur).your
Thalassomedon
Vert.
Paleo.
haningtom
(long-necked
I5(suppl. to
5)161
A.
University Press.
Wiman, C. J. 1 9 2 0 . Some reptiles from the Niobrara
group in Kansas.
Sci.
Acad.
T h e first record of
27:72627.
Geol.
Water
Acad.
Jour.
Clobidens
Alabama
Elasmosaurus,
21:22136.
Jour.
Nat.
Alabama.
discovered at W h i t b y in February
22:565.
Cimoliasaurus,
A new specimen of
1985.
world's
12:4551.
Geol.
the
4 5 6 : 1 51.
2:177.
Pub.
mosasaurs.
Geology.
osborni,
1(9)241-44.
1905.
World:
Vert.
Modern
the
Kansas,
4(5)81547.
1902.
of
Changed
Stupendemysgeographicus,
Breviora
from
1988.
russelli,
Paleontology
that
Mosasaurs.
1898.
Birth
1976.
States, your
4 ( 4 > 7 7 - 8 5 .
the
Map
5 6 ( 5 ) 1 0 2 (abstract).
2 8 0 5 ) 5 0 .
C.
alabamaensis
1894.
R.
Wright, K. R.
2(2)8584.
Quart.
and
largest turtle.
18(465)545.
Science
Smith
The
HarperCollins.
Wood,
i6(4o5):262.
2001.
Bull.
Geol.
Instn.
Uppsala
18:918.
Bull.
Inst.
Egypte
Index
Academy o f Natural Sciences, Philadelphia, 9 4 , 1 2 7 -
jaws, 156
129
Amniotes, 2 7 , 2 8 , 2 9 , 8 5 n
28
Acanthostega,
Amphibians
Adams, Dawn, 1 8 9
breathing of, 50
Adetatde,
Aegirosaurus
as index fossils, 12
eggs, 2 9
174
Anapsids, 2 8 , 8 5 n
9697
leptospondylus,
Ando, T., 7 8 7 9
Africa
fossils found in, 1 7 4 , 1 8 7 , 2 2 2 2 2 5
human fossils. 10
Aigialosaurs,
200202,
2oi(illus.),
Augolasaurus
225
bocagei,
Anguid lizards, 2 4 9 2 5 0
Aigialesaunts
250
258
Ankylosaurus,
200
novaki,
(slowworm),
AnguisJragilis
200
dahnaticus,
Anning, Joseph, 6 7 , 6 9
Alabama
fossils found in, 2 1 5 , 2 1 6 2 1 7 , 2 1 9 2 2 0 , 2 2 8
Anning, Mary, 3, 2 1 , 6 7 , 6 9 7 0 , 1 0 6 ,
Selma Formation, 2 1 5 , 2 1 9 , 2 2 0
1-4
Anning. Richard, 6 7
Antarctica, 2 1 5 , 2 1 7 , 2 6 1 2 6 2
Albino, A . M., 2 4 7
Apatosaurus,
Alexander, Annie, 8 8 , 9 3 , 1 2 2
Alligator
Allosaurus,
lizards
1,
(Gerrbenotus),
Archaeonectrus,
250
255
rostratus,
Archaeopleryx,
26,
42,
120,
95,
Archosaurs, 2 9
Argentina
192
140,
258
Alzadasaurus
pcnibcrteni,
258
168
Alvarez, Luis, 2 5 3 , 2 5 4
kansasettsis,
155a
turtles, 32
130
130,
157158
Aristotle, 1 7 , 18
Ammonites
Aspect ratios ( A R ) , 1 4 2 1 4 3
leathcrback
turtle
(Dermochelys
coriacea),
33,
Attenborosaurus,
161,
Bernard, L., 2 2 1
Bible, 1 7 , 18
53(illus.)
Billfish, 4 8
i62(illus.)
Birch, Thomas, 6 7 , 6 9
Australia
Birds
ggs. 49
endothcrmic nature, 4 5
172175,
i73(illus.)
pliosaur fossils, 1 7 5 , 1 7 6
fishing by,
I53n
gizzards, 156
Australian Museum, 1 7 3
Avascular necrosis, 2 3 9
Blainvillc's
Axial oscillation, 7 9
Blake, J. R, 1 6 6
beaked
whales
(Mesoplodon
densiroslris),
Blake, R . W , 1 0 .
Baily, W. H.. . 7 .
Blucfin tuna, 6 2 ( i l l u s . )
1 5 2 , 1 6 1 , 163, 1 6 4 , 188, 1 8 9 1 9 0
Baleen whales, 5 7 , 9 3
Baptancdon.
See
Ophtbalmosaurus
Bardet, Nathalie, 9 6 9 7 , 1 1 4 , 1 6 1 1 6 3 , 1 8 7 , 2 1 8 ,
220
Basilosaurus,
Bones
analysis of, 5 2 , 2 3 4
descriptions of, 5, 7
low density, 2 0 5
12
Barosaurus,
Bonde, Niels, 2 2 5 2 2 7
125
pachyostosis, 1 9 1 , 2 4 0 , 2 4 7
terminology, 5
Baur, Gcorg, 8 4 , 85
Bottlenosc dolphins, 6 2 ( i l l u s . )
BBC,
Brachauchenudae,
179
Bclemnites, 1 0 0 , 2 1 5 , 2 i 5 n
Braehauchemus,
Belgium
Braebiosaurus,
180,
192
i83(illus.),
elasmosaur fossils, 1 6 3 , 2 6 2
Braehypterygius,
2 2 1 , 230231
183184,
192,
193
itfn
109
British Columbia
Mons basin, 2 2 0 2 2 1
Burgess Shale, 9
mosasaur fossils, 7 , 2 4 , 1 9 5 1 9 8 , 2 1 8 , 2 2 0 2 2 1 , 2 2 9 ,
Bends, i n 1 1 2 , 2 3 9 , 2 4 0 2 4 1
Brontosaurus,
Bcnnison, Alan, 2 1 4
Broom, Robert, 2 2 2
258
24on
Centropherodes
Buchcr, Hugo, 82
Cetaceans
(dogfish
appendiculatus
sharks),
Buchy, Marie-Celine, 9 5 , 1 8 1 1 8 2
blubber, 5 2 5 3 , 5 4
Buckland, Frank, 1 1 9
bone densities, 2 4 0
Buckland, William, 1 9 2 0 , 6 7 , 7 0 , 1 1 9 , 1 2 4
evolution of, 5 7 5 8 , 1 0 5 , 1 9 9
Bud, J. W ,
250
"Bunker"
reproduction,
Tylosaurus,
216
See also
Caldwell, Michael W , 1 4 , 4 0 , 5 6 5 7 , 1 0 0 1 0 m , 2 1 6 ,
217218,
234,
244,
245248,
249
(Zalophus),
63,
Cbaobusaurus,
80,
mydas
(green turtle),
80,
Chensaurus,
81
80
geishanensis,
33
104
89
Callawayia,
95
Cbacaisaurus,
Cbeloma
232
30, 73
Dolphins; W h a l e s
China
130
Callawaysaurus,
Camp, Charles, 9 0 9 2 , 2 1 4
dinosaur fossils, 95
iehthyosaur fossils, 8 0 , 9 3 9 4 , 9 5 9 6
Yixian Formation, 9
Camper, Pieter, 1 9 5 1 9 6
Canada, fossils found in, 9 , 8 9 9 0 , 2 1 6 , 2 3 0
Chonglakmani, Chongpan, 8 0
Carcbarodon
Christiansen, Per, 2 2 5 2 2 7
carina
(loggerhead turtle),
33,
34(illus.)
CIcma, John, 1 7 4
Ciidastes,
Carinodens
221,
belgicus,
229
lengths, 2 0 4
Carpenter, Simon, 1 6 9
predators, 2 2 8
Carrier, D. R., 49
propython,
skulls, 2 0 5
sternbergi,
217218
Carsosaurus,
marchesetti,
200
Carte, Alexander, 1 7 1
Casiiuria
kiddi,
28
Catsharks, 7 8 7 9
Caypullisaurus,
205(illus.),
204206,
210
bone densities, 2 0 5 2 0 6
fossils, 2 0 3 , 2 2 0 , 2 2 8 , 2 4 2
229
fraasi,
35
Claudiosaurus,
264
megalodon,
Caretta
Chronological periods, 1 1 1 3
79
carcharias,
95
219
213,
214
tail fins, 2 0 4 2 0 5
water depths, 2 1 9 , 2 2 0 , 2 3 9
Coates, Michael I., 2 4 9
Colagrande, John, 1 4 7 1 4 8 , 1 5 2 , 235
Colbert, E. H., 63
Coles, Henry, 9 7
218
C r u i c k s h a n k , A r t h u r R . I.,
132,
trochanteric,
Coniasaurus,
crassidens,
172,
136
16,
Cryptoclidus,
40
eurymerus,
40
oxoniensis,
87
Contectopalatus,
176,
186,
I34(illus.),
132133,
132,
133,
134135, 1 6 7 1 6 8 , 1 6 9 1 7 1 ,
191
I33(illus.),
172
134,
184
133
132
ricbardsoni,
Convergent evolution, 4 0 , 4 8 , 6 1 , 9 3
Cumbaa, S. L., 2 1 6 , 2 3 4
Conybeare, W i l l i a m Daniel, 7 0
on ichthyosaurs, 71
Curry, Phil, 2 2 6
o n mosasaurs, 2 4 , 1 9 7
Custer, George, 2 1 2
plesiosaur name, 1 9 , 1 1 8
o n plesiosaurs, 6 6 , 6 7 , 6 9 , 1 1 9 , 1 2 3 , 1 2 4 , 1 2 5 , 1 2 6 ,
Cymbospondylus,
9394,
1J7
94(illus.),
104
94
asiaticus,
Darwin, Robert, 1 9
128
description
of
Elasmosaurus
56
platyurus,
o n mosasaurs, 2 1 0 , 2 1 9 , 2 4 5
Deeming, D. G, 4 9 , 7 3 7 4
(dorado,
mahi-mahi),
239
Delair, Justin B 67
Dermochelys
Cragin, F 1 8 9
32,
5354,
Diapsids, 2 8 2 9 , 2 0 4
247
Dinilysia,
Cretaceous era, 1 1 1 2
shark),
241242
Dinosaurs
American Museum of Natural History exhibits,
Crocodiles
12
ballast stones, 1 5 7
cctothermic nature, 4 5
marine, 4 4 , 5 8 , 1 0 4
diets, 4 2
saltwater, 58, 2 1 6
Crocodylus
porosus
58
Creationists, 6 3 6 4
(lamnid
(leatherback turtle),
mantelli
255
Cowen, Richard, 2 9 , 4 9 5 1 , 1 0 0 , 2 3 3
Cretoxyrhina
46,
Dcinonychus,
Cope's rule, 1 9 7 , i 9 7 n
hippurus
Dawson, J . W , 2 8 , 4 2 , 165
DcBraga, Michael, 2 0 0 , 2 0 1
mosasaur names, 2 0 8 , 2 0 9 , 2 1 0
Coryphaena
(saltwater crocodile),
Crombie, Andrew, 1 7 6
58,
216
endothermic, 4 6 4 7
extinctions, 1 4 , 15, 1 6 3 , 2 5 3 2 6 0
feathered,
95
192
Edgarosaurus,
fossils, 9 , 1 9 2 0 , 2 6 , 4 6 , 1 7 0 1 7 1
growth patterns, 4 7
limbs, 41
Ein Yabrud, 2 4 6
Elasmosaurs
name, 2 0 , 25
attacked
by
Kronosaurus,
178179
scientific names, 4
study of, 2 5 2 6
distribution,
161,
extinctions,
161
weights, i 8 2 n
feeding behavior, 1 4 8
Diplodocus,
42,
fossils, 9 1 0 , 1 3 0 , 1 3 2 , 1 5 7 1 5 8 , 1 6 3 1 6 4 , 1 7 8 1 7 9 ,
!55n
Dogfish sharks, 2 1 7 , 2 1 8
189,
Doliehorhynehops,
osborni,
190,
257
192
hunting behavior, 1 5 4
130
Dollo, Louis, 1 9 8 , 2 1 1 , 2 1 4 , 2 2 1 , 2 2 9
Dollosaurus.
See
necks, 1 2 1 1 2 2 , 1 5 4 1 5 5 , 1 7 8 1 7 9
reproduction,
Prognathodon
Dolphins
iVc
beaching,18
also
147148
Plesiosaurs
Elasmosaurus
bottlenose, 6 z ( i l l u s . )
morgani,
ccholocation, 9 8
platyurus,
159160
127129,
flippers, 14
Elephants, 2 4 , i 8 2 n
hunting behavior, 2 ( 9
Enaliosaurs,
leaping, 101
England
locomotion, 4 1 ,
103
62(illus.),
I28(illus.),
130,
153
166
dinosaur fossils, 1 9 2 0
164
7 m , 10010m
fossil sites, 9
ichthyosaur fossils, 6 5 6 6 , 6 7 7 3 , 8 1 8 2 , 1 0 5 1 0 7
tail fins, 1 4 1 5
viviparous reproduction, (o
Dorado
(Ceryphaena
hippurus),
239
Dortangs, Ruud W 2 2 1 2 2 2
Douglas, Kate, 6 4 6 5
I )o\ le, Peter, 1 0 0
plcsiosaur fossils, 6 9 , 1 2 0 , 1 2 3 1 2 4 , 1 2 5 1 2 7 , 1 3 2
133, 135, 1 5 0 , 1 5 7 , 1 6 0 1 6 1 , 1 7 4
pliosaur fossils, 1 6 6 1 6 8 , 1 6 9 , 1 7 1 1 7 2 , 1 8 6 , 1 8 7 ,
191
Enzymes, 5 6 5 7
Durophagy, 8 7 , 2 2 9
Eretmochelys
imbricata
(hawksbill
turtle),
33,
33(illus.)
Erickson, G . M., 4 7
Ectenosaurus,
215,
217
172174,
i73(illus.)
81,
Eurhinosaurus,
Figuier, Louis, 2 2
82
io7(illus.),
longirostris,
107109
ectothermic nature, 4 8
167168
arcuatus,
Fishes
circulatory systems, 4 8
192
Eurycleidus,
eggs, 2 9
fossils, 2 1 8
241242
hunting behavior, 2 3 9
Evolution
leaping, 101
convergent, 4 0 , 4 8 , 6 1 , 9 3
locomotion, 4 1 ,
creationist view, 6 3 6 4
swordfish, 103, 1 0 7 , 1 0 8
evidence of, 64
extinctions and, 25
82,
Excalibosaurus,
106107
Extinctions
caused b y asteroid (K-T extinction), 15, 4 4 , 2 0 4 ,
136
turtle
(Nalator
depressus),
34
Darwinian explanations, 2 3 , 2 5 , 2 6 0
descriptions of, 56
dinosaur, 9 , 1 9 2 0 , 2 6 , 4 6 , 1 7 0 1 7 1
of elasmosaurs, 161
early discoveries, 1 8 2 1 , 2 2
evidence of, 2 4 2 5
explanations of, 2 3 , 2 5 3 2 5 5 , 2 5 9 2 6 0
of ichthyosaurs, 1 4 , 4 4 , 6 4 , 1 1 4 1 1 6 , 2 0 0 , 2 5 7
human, 1 0 , 25
of marine reptiles, 1 6 1 1 6 2 , 2 5 7 2 6 0
incompleteness of record, 89
of mosasaurs, 1 4 , 2 0 3 , 2 0 4 , 2 4 1 , 2 4 2 2 4 4 , 2 5 7
index, 12
of plesiosaurs, 1 4 , 4 4 , 1 2 1 , 1 6 1 1 6 4 , 2 5 7
of marine reptiles, 2 0 2 1 , 2 2
recent, 2 6 0
Eyes
study of, 7 8 , 9 1 1
See
also
specific
Fraas, Oskar, 7 4
of plesiosaurs, 135, 1 6 0 1 6 1
France
o f pliosaurs, 1 6 9 , 1 7 2
See also
Sclerotic rings
marine
reptiles
ichthyosaur fossils, 82
pliosaur fossils, 1 8 2
Frcy, Ebcrhard ("Dino"), 9 5 , 139, 1 4 4 , 1 4 6 , 1 8 1 1 8 2
Gallomimus,
Gaskill, P., 36
Field Museum, 2 1 9
Gastralia, 1 4 0 1 4 2
255
213,
214,
sternbirgi,
40
221,
225
213214
platyspondylus,
fossils,
Henodus
211,
Halisaurus,
197
Geosaurus,
230
pembinensis,
219
Holzmaden, 9 , 7 4 7 5 , 8 1 , 8 6
ichthyosaur fossils, 7 4 - 7 5 , 8 1 , 8 6 , 9 6 9 7 , 1 1 4
155,
pistosaur skeleton, 1 1 9 1 2 0
Solnhofen, 9 , 9 6 9 7 , 2 0 2
Hampe, Oliver, 1 7 7
(alligator lizard),
Cerrhonotus
1 8 1 , 1 8 5 1 8 6 . See also
Halvorson, Dennis, 2 4 2
Hanson, Mike, 2 4 2
250
Harlan, Richard, 4 2 , 1 2 4 1 2 5 , 1 9 8
Gibbes, Robert, 2 1 9
Harp
Gilmore, Charles W 2 0 6 , 2 1 9 , 2 2 8
Ginsu sharks, 2 4 1 2 4 2
seals
(Phoea
111
groenlandica),
Zoology, 1 7 5 1 7 6
Hawkins, T h o m a s W., 2 1 , 7 0 7 1 , 1 6 0
Gizzards, 1 5 6 , 157
Hawkins, Waterhouse, 2 5 2 6
Glass
snakes
Globidens,
216,
221,
Hawksbill
250
(Ophisaurus),
219,
228(illus.),
227,
Henodus,
Godfrey, S. J 233
217,
Heterodontus
40
(horn
shark),
Great Flood, 1 8 , 23
Holmes, R., 2 1 4 , 2 1 6 , 2 3 4
229
Hoganson, John, 2 4 2
Gould, Stephen J 1 3 1 4 , 7 m
(Carcbarodon
earebarias),
59,
79,
216,
Holzmaden, Germany, 9 , 7 4 7 5 , 8 1 , 8 6
Home, Sir Everard, 6 7 6 8
Horn
Green, Matthew, 1 2 6 1 2 7
Green
turtle
(Cbeloma
mydas),
Grippia,
(Heterodontus),
229
Hotton, Nicholas, 2 5 3
Hovasaurus,
109
mordax,
sharks
33
Gregory, Joseph, 92
Crendelius
35,
36(illus.)
Hua, Stephane, 1 8 7
80
Haasiopbis
terrasanctus,
246,
Hunting behaviors
247
ambush prcdation, 8 0 , 1 2 0
Haines, T i m , 1 7 9 , 1 8 0 1 8 1
Hainosaurus,
bemardi,
229230,
221,
226,
33,
HofFmann, C. K., 1 9 5 , 1 9 7
224(illus.)
223224,
222223
nigeriensis,
imbricata),
4o(illus.)
3840,
cbelyops,
Goldfuss, August, 1 9 8
Goronyosaurus,
(Eretmochelys
Heatwole, H., 2 4 5 , 2 4 6
219
alabamaensis,
turtle
Hayden, Frederick, 2 0 7
228-229
23i(illus.),
227,
230231
257
o f dolphins, 2 3 9
of elasmosaurs, 1 5 4
33(illus.)
Hunting
behaviors,
84
Macgowama,
continued
o f ichthyosaurs, 8 0 , 9 9
Macroplerygius,
of mosasaurs, 2 3 5 , 2 3 9 2 4 0
Mixosaurus,
of plesiosaurs, 1 2 0 , 1 4 3 , 1 5 2 1 5 4
Mollesaurus,
96
Omphalosaurus,
of sharks, 2 3 9
Pbalaradon,
80,
85,
29
88(illus.),
Sbaslasaurus,
Shonisaurus,
Ichthyopterygia, 8 4 , 8 8
Stenopterygius,
Iehthyosaur fossils
Suevoleviathan,
44,
64,
Svalbardosaurus,
in Asia, 7 8 7 9 , 8 0 8 1 , 9 3 9 4 , 9 5 9 6
Temnedontosaurus,
in Australia, 9 9 , 1 1 3
Tbaisaurus
ToretocnemttS,
in E n g l a n d , 6 5 6 6 , 6 7 7 3 , 8 1 8 2 , 1 0 5 1 0 7
Utatsusaurus,
fetuses, 7 2 7 4 , 81
Wimanius,
in France, 82
See
63,
80,
80,
Crippia,
Isfjordosaurus,
Leptonectes,
105106,
io6(illus.),
82,
82,
94(illus.),
io7(iIIus.),
106107
78(illus.),
7879,
8081
86
also
Ichthyosaurus;
Ophthalmosaurus
Ichthyosaurs
104
107109
earliest, 7 8 8 0
109
extinctions, 1 4 , 4 4 . 6 4 , 1 1 4 1 1 6 , 2 0 0 , 2 6 1
80
eyes, 7 7 , 9 6 , 9 9 , 1 0 5 , 1 0 9 1 1 1
84
8182,
109
96
diets, 1 6 , 8 7 , 9 8 , 9 9 , 1 0 0 , 1 0 5 1 0 6 , 1 1 2 , 113
87
81,
103
80
descriptions of, 66
9394,
Crcndelius,
81,
81
104
Contectopalatus,
txcaltbosaurus,
68,
colors, 9 9 1 0 0
95
Eurhinosaurus,
104
8}(illus.),
chronology, 7 5 7 7
95
Cymbospondylus,
82,
breathing of, 6 3 , 9 9 , 1 0 0 , 1 0 2
89
Caypullisaurus,
Chensaurus,
81,
9i
93,
brain sizes, 9 7
109
Chaohusaurus,
75,
chonglahnanii,
64, 7980,
9697
Chacaisaurus,
9i(illus.),
2 _
80
body plans,
Iehthyosaur genera
Callawayia,
74,
aquatic adaptation, 4 3 , 6 4 6 5
in New Zealand, 2 2 7
Brachypterygius,
8992,
8889,
86
in Argentina, 95
in Germany, 7 4 7 5 , 8 1 , 8 6 , 9 6 9 7 , 1 1 4
113114,134
96
Quamcbtbyosaurus,
28,
ii2(illus.),
96
Qianiebthyosaurus,
25
Aegirosaurus,
87
8 6 - 8 7
99,105,
Platyptetygius,
122
alexandrae,
Hylaeosaurus,
Hylonomus,
130
scrpenlinus,
Hydrotherosaurus
8587
95
of pliosaurs, 1 7 2 , 1 8 0 1 8 1 , 1 9 1
Hydrahnosaurus
77(illus.),
7677,
106107
flippers, 1 4 , 1 1 4
62(illus.
fore-flipper bones, 1 0 5 , i o 5 ( i l l u s . )
Index fossils, 12
gastralia, 1 4 0 , 141
hunting behavior, 8 0 , 9 9
215, 2 2 0 2 2 1 , 230231
largest, 9 0 9 2 , 9 3 9 5
Isfjordosaums,
leaping, 1 0 0 1 0 2
Israel
lengths, 4 4 , 1 0 4 , 2 2 o n
limbs, 4 1 , 6 5 , 7 6 , 1 1 4 , 134
in literature, 2 1 2 2
84
locomotion, 8 , 4 2 , 5 0 5 2 , 7 9 , 1 0 2 1 0 3 , 1 0 4
origins, 6 3 6 4
Jakobsen, Sten, 2 2 5
ribs, 6 3 , 8 6
Japan
62(illus.),
7 m , 10010m
ichthyosaur fossils, 7 8 7 9 , 8 0 8 1
similarities to sharks, 11
Journey
skin, 8 , 9 7
Jurassic era, 11
to
the
Center
of the
Earth
(Verne),
2122
skulls, 4 3 , 8 7 , 9 7 9 9
swordfish, 1 0 7 1 0 9
Kaiwhekca
Kansas
134135
kaitiki,
taxonomy, 7 5 , 8 3 8 7 , 9 3
Benton Formation, 1 8 9
teeth, 8 7 , 9 8
mosasaur fossils, 1 9 9 , 2 0 3 , 2 0 6 , 2 0 7 2 0 8 , 2 0 9 2 1 0 ,
toothless, 1 1 2 1 1 3
plesiosaur fossils, 1 2 7 , 1 5 7 , 1 5 8 1 5 9
vertebrae, 7 8 , 7 9
pliosaur fossils, 1 8 3 1 8 4 , 1 8 8 1 8 9
viviparous reproduction, 4 3 , 4 8 4 9 , 7 2 7 4 , 7 6 ,
97-98,
99
See also
Marine reptiles
See also
44,
Ichthyosaurus,
68.
72(illus.)
105
hreviceps,
communis,
71,
73,
107
longirostris,
Iguanodon,
96
leptospondylus,
Kimmerosaurus,
28
20,
25,
136
198
missouriensis,
(Lepidochelys
26,
kempii),
Kesling, Robert V , 2 3 5 2 3 6 , 2 3 7 2 3 8
81
cornaUamts, 85
Ichthyostega,
85
atavus,
Kase, T., 2 3 7 , 2 3 8
255
Knight, Wilbur, 1 8 8
34
214
Kolposaurus,
Leptopterygius
214
bennisoni,
dragon
(Varanus
175179,
Libonectes
180,
185,
192,
176177,
I77(illus.)
238,
Liodon,
196197
Leatherback turtles
(Dermochelys),
32,
Michael
S.
Y.,
217218,
33,
245247,
53(illus.),
See
210,
See
also
248249
2o8n
Leiodon
I78(illus.),
179,
rossicus,
181
skeleton, i 8 i ( i l l u s . )
skulls, 1 7 9 , 1 8 4 , 193
Lepidocbelys
tail fins, 1 4 2
teeth, 4 3 , 1 7 9 , 1 8 0 , 1 8 4
34
34
Lithophagic animals, 1 5 8
Lepidosaurs, 2 9
Leptoclcididae,
LeptocUidus,
elemai,
192
I73(illus.),
172175,
Leptonectes,
moorei,
tenuirostris,
200202,
2oi(illus.),
anguid, 2 4 9 2 5 0
174
ectothcrmic nature, 45
8182
monitor, 4 8 4 9 , 2 3 9
68
aeutirostris,
Lizards
alligator, 2 5 0
174
superstes,
192
aigialosaurs,
174
capensis,
192
lengths, 1 7 9
234235
Liodon
olivaeea,
185,
182
Lemonnier, A., 2 2 1
kempii,
179182,
diets, 183, 1 8 4
234
also
53
Jerox,
Leidy, Joseph, 2 1 1 , 2 1 9
mosasauroides,
209
207,
Liopleurodon,
Leiodon,
207208
proriger,
79
54- 58
Lee,
250
nepaeelicus,
nasus
210,
dyspelor,
242244
gigantea,
159160
2 2 0 , 2 2 1 , 2 2 2 , 2 2 3 2 2 5 , 2 2 9 , 230233, 234235,
177178
queenslandicus,
130,
nwrgani,
193
attacks on elasmosaurs, 1 3 2
Lacerta
Leptonectes;
Li, Chun, 9 6
Kourlis, George, 9 9
boyacensis,
also
Lhwyd, Edward, 6 5 6 6
58
komodoensis),
Kosch, Bradley, 92
Kronosaurus,
See
Ley, Willy, 7 4
214
tuckeri,
Komodo
86.
disinteger,
Temnedontosaurus
similarities to mosasaurs, 11
82
82,
106107
Leptonectidae, 8 2
varanid, 11
See
also
Reptiles
214,
217218,
242
Macroplerygius
Locomotion
96
posthumus,
aquatic, 4 8
96
trigonus,
axial oscillation, 7 9
Macrosaurus
208,
proriger,
2o8n
of cetaceans, 4 1 , 1 0 3 , 136
Mahi-mahi
drag and, 5 1 5 2
of fishes, 4 1 , 136
M a k o sharks, 7 9
of ichthyosaurs, 8 , 4 2 , 5 0 5 2 , 7 9 , 1 0 2 1 0 3 , 1 0 4
Mamenchisaurus,
methods, 1 3 6 1 3 7
Mammals
(Coryphaeua
hippurus),
I22n
of mosasaurs, 15, 4 1 , 2 0 4 , 2 1 6 , 2 2 5 , 2 3 1 2 3 4
a s amniotes, 2 7 , 2 8
o f penguins, 1 3 7 , 138, 1 3 9 , 1 4 0 , 1 4 5 , 1 9 9
endothcrmic nature, 4 5
of plesiosaurs, 1 3 4 , 1 3 6 1 4 7 , 1 4 8 1 4 9 , 1 5 1 1 5 2 , I58n,
239
largest, i 8 2 n
marine, 5 2 5 3 , 5 4
175
of pliosaurs, 15, 1 4 4 , 1 4 8 1 4 9 , 1 8 2 1 8 3 , 1 8 9
ribs, 6 ; n
viviparous reproduction, 30, 4 9
of sea snakes, 1 3 6
of sea turtles, 1 3 7 , 138, 1 3 9 , 1 4 0
Manabe, Makoto, 7 3 7 4 , 8 9 , 9 6
of sharks, 1 0 2 , 1 0 3 , 136
Manatees,
of sircnians, 136
of tuna, 103
Mantell, Gideon, 2 0 , 2 6 , 7 0 , 1 1 9
underwater flight, 1 6 , i o 2 n , 1 3 7 , 1 3 8 , 1 4 4 , 2 3 2 2 3 4
Mantcll, M a r y Ann, 2 0
of whales, 41
Maresaurus,
caretta),
33,
34(illus.)
coecai,
1 9 1 . See also
Sircnians
192
180,
187188
Marine reptiles
Londish, Sid, 1 7 3
London Brick Company, 137138
aquatic adaptation, 5 8 , 1 9 9 2 0 0
Long, ] . , 1 8 6 , 2 4 5
blubber, 53
Long, J . A., 1 7 6 , 1 8 6 , 2 0 3 , 2 4 5
colors, 8
Coniasaurus,
current species, 58
diets, 55
Lyell, Charles, 1 9 , 4 2
40
longirostris,
tenuiceps,
extinctions, 1 6 1 1 6 2 , 2 5 7
187
Machimosaurus,
Maeroplata,
evolution of, 3 0 , 57
84
Macgowania,
166,
energy needs, 5 4 5 5
Henodus,
192
4o(illus.)
largest, 1 7 6 , 1 7 9 , 2 2 0 , 2 2 o n
166
166167,
3840,
i67(illus.)
in literature, 2 1 2 2
Marine
reptiles,
32
Meiolania,
continued
metabolisms, 2 9 , 4 6 , 4 7 , 4 8 5 1 , 5 6 5 7 , 1 0 4
Mesophis
nopscai,
247
Mesoplodon
densirostris
(Blainvillc's
pistosaurs, 1 1 9 1 2 0
Mesosaurus,
30,
placodonts,
3738,
39(iIIus.),
3i(illus.),
beaked
whale),
2 on
4
57
Mesozoic era, 11
41
Metabolism
reproduction, 1 4 , 2 9 3 0 , 4 8 4 9 , 5 8
of dinosaurs, 4 6 4 7
sauroptcrygians, 3 4 3 6
cctothcrmic, 2 7 2 8 , 2 9 , 4 5 , 52
scientific names, 4 5
endothermic, 2 9 , 4 5 , 4 6 4 7 , 4 8 5 1 , 5 6 5 7 , 1 0 3 1 0 4
skulls, 4 3
enzymes and, 5 6 5 7
variations among, 16
of fishes, 48
of ichthyosaurs, 1 0 3 1 0 4
See also
Pliosaurs
of marine reptiles, 4 5 , 4 6 , 4 7 , 4 8 5 1 , 5 5 5 7 , 1 0 4
of mosasaurs, 2 3 8 2 3 9
of reptiles, 2 7 2 8 , 2 9 , 4 5 , 52
of sea turtles, 54
assistants, 8 4
fossils collected by, 2 0 7 , 2 1 1 2 1 3 , 2 5 0
Metashastasaurus.
mosasaur names, 2 0 8 , 2 1 3 2 1 4
Minoura, N 7 8 7 9
See
Callawayia
Minton, S . A., 2 4 5 , 2 4 6
Mixosauridac, 85
Martin, John, 21
Mixosaurus,
7677,
M a r t i n , L. D., 1 5 8 1 5 9 , 2 0 3 , 2 0 5 , 2 3 9 , 2 4 0
atavus,
comalianus,
232,
238-239
major,
maotaiensis,
nutans, 85
Mazin, J.-M., 8 0
nordeskioeldii,
99-
> 103
Moanasaurus
7 6 , 8586
8687
McGowan, Chris, 4 7 , 4 8 , 6 9 7 0 , 7 4 , 7 5 7 6 , 8 1 , 8 2 ,
77(illus.)
86
85
85
mangahouangae,
227
Moisley, W 10
Mollesaurus,
95
Monitor lizards
McNamara, K 1 8 6 , 2 4 5
Monster of Aramberri, 1 8 2
Megalama
prisca
Megalneusaurus,
Megalosaurus,
(monitor lizard),
180,
20,
188,
239
prisca),
Montana
Bear Gulch Formation, 9
192
dinosaur fossils, 2 6 , 4 6
25
3 0 2 S' A 'D %A g O T V
5859,
(Megalama
5859,
239
bone densities, 2 0 5 , 2 4 0
I loll ( rook I 01 i n a l i o n , y
deep divers, 2 3 9 2 4 1
Moore, Chris, 82
diets, 1 6 , 1 9 8 , 2 1 5 , 2 2 3 , 2 2 6 , 2 2 8 2 3 0 , 2 3 4 2 3 8
distribution,
Mosasaur fossils, 2 0 2 2 0 3
earliest, 1 9 9 , 2 0 3
215
in Belgium, 7 , 2 4 , 1 9 5 - 1 9 8 , 2 1 8 , 2 2 0 2 2 1 , 2 2 9 , 2 5 1
evolution of, 2 0 2 2 0 3 , 2 1 4 , 2 4 4
extinctions, 1 4 , 2 0 3 , 2 0 4 , 2 4 1 , 2 4 2 2 4 4 , 2 5 7
embryos, 2 1 7
flippers, 2 0 4
hunting, 235, 2 3 9 2 4 0
in Kansas, 1 9 9 , 2 0 3 , 2 0 6 , 2 0 7 2 0 8 , 2 0 9 2 1 0 , 2 1 1 ,
largest, 1 9 7
latest species, 2 4 4 , 2 5 7
213, 215, 2 1 6 , 2 1 8 2 1 9 , 2 4 1 2 4 2 , 2 5 0
locations, 1 9 9 , 2 0 3 , 2 1 1 , 251
lengths, 1 9 7 , 2 0 3 , 2 0 4 , 2 1 0 , 2 1 6 , 2 2 3 , 2 2 6 , 2 3 0
young, 2 1 8
locomotion, 15, 4 1 , 2 0 4 , 2 1 6 , 2 2 5 , 2 3 1 2 3 4
metabolism, 2 3 8 2 3 9
Mosasaur genera
221,
Carinodens,
possible ancestors, 2 0 0 2 0 2
229
215,
Ectenosaurus,
Clohidens,
name, 1 9 7
225
Angolasaurus,
217
l i b ,
228(illus.),
GoroHyosnMnu, 2 1 7 , 2 2 2 2 2 4 ,
Hainesaurus,
221,
226,
Hahsaurus,
211,
213214,
Kolposaurus,
Leiodon,
210,
227,
221,
23i(illus.),
257
reproduction, 1 9 9 , 2 1 7 , 2 1 8
ribs, 1 4 1
225
214
scales, 8
234235
shallow-water,
skulls, 2 0 2 , 2 0 4 , 2 0 5 , 2 1 6 , 2 1 8 , 2 2 3 , 2 3 0
2o8(illus.)
208,
219
shark attacks. 2 1 7 . 2 2 1 - 2 2 2 . 2 4 1 - 2 4 2
Liodon, 2 0 7 2 0 8 , 2 o 8 n , 2 0 9 , 2 1 0 , 2 5 0
Macrosaurus,
250
replacement of ichthyosaurs, 2 0 0
224(illus.)
229231,
219,
228229
Moanasaurus,
227
snouts, 2 0 8 , 2 3 0
Oronosaurus,
226
tails, 2 0 4 , 233
taxonomy, 2 o 8 n , 2 1 3 2 1 4 , 2 2 7
214
Phospborosaurus,
teeth, 1 9 7 , 1 9 8 1 9 9 , 2 1 5 , 2 2 3 , 2 2 4 , 2 2 6 , 2 2 8 2 2 9 ,
214
Plotosaurus,
224225
234-238
Rikkisaurus,
227
time period, 13
Selmasaurus,
217
vertebrae, 2 1 8 , 2 3 0 , 2 3 1 2 3 2
Pluridens,
See
also
vision, 1 9 7 1 9 8
227
Tantwhasaurus,
Cltdastes;
Prognathodon;
Mosasaurus;
Platecarpus;
Tylosaurus
Mosasaurs
Plioplatecarpus;
weights, 2 1 6
See also
Mosasaurus,
Marine reptiles
197,
aquatic adaptation, 1 9 9 2 0 0 , 2 1 4
conodon,
zzi
avascular necrosis, 2 3 9
copeanus,
211
203,
210,
214,
220,
225
J NT) EX
303
continued
ichthyosaur fossils, 2 2 7
mosasaur fossils, 2 0 3 , 2 2 7 2 2 8 , 2 4 4
Mosasaurus,
boffmanni,
2 3 0 2 3 1 , 251
pliosaur fossils, 9 1 0
216
horridus,
221
lemonnieri,
198
maxitniliani,
198,
missouriensis,
Niger, 2 2 4 2 2 5
216
Nigeria, 2 2 2 2 2 3
227
mokoroa,
nigeriensis,
zzz
in,
plesiosaur fossils, 1 2 9 1 3 0
112113
Mullcr, Simeon, 9 0
beloclis,
leedsii,
132,
132,
Noe,
L,
Nolf,
135
135,
32
Niolamia,
i22(illus.)
121,
Muraenosaurus,
mosasaur fossils, 2 1 1 , 2 1 3 , 2 1 5 , 2 1 8
180,181,182-183,186-187,
D,
'9'
.98
Norman, D. B 1 7 0
184
depressus
(flatback
turtle),
34
36,
41,
140
34
tchernovi,
3;(illus.),
34,
Notkosaurus,
Novacek, Michael ] . , 2 5 6
O'Fallon, Benjamin, 1 9 8
O'Keefe, F. R., 3 6 , 1 4 2 1 4 3 , 1 7 4
Omphalosaurus,
80,
nevadanus,
87
Necks
87
of dinosaurs, i 2 2 n , i 5 5 n
of elasmosaurs, 1 2 1 1 2 2 , 1 5 4 1 5 5 , 1 7 8 1 7 9
Opetiosaurus,
of plesiosaurs, 1 1 8 , 1 2 0 , 1 2 1 1 2 2 , 135, 1 5 3 1 5 5 , 1 6 0 , 1 6 8
of pliosaurs, 1 8 3 , 1 8 6
mosasaur fossils,
Ophisaurus
7, 2 4 , 195198,
I97n,
218, 220
2 2 1 , 2 2 9 , 251
Teylers Museum, 2 1 5
I73(illus.)
200
huccichi,
Ophiomorphus,
Netherlands
172175,
242
246
(glass snake),
Ophthalmosaurus,
250
97(illus.),
no(illus.)
Ncubig, Bcrnd, 9 2
edentulousness, 1 1 2 1 1 3
eyes, 109no, in
fossils, 95
Newton, Isaac, 65
New Zealand
elasmosaur fossils, 9 1 0
swimming speeds, 1 0 4
Orndorff, R . L , 9 1
Penguins
226
Oronosaurus,
Osborne, Roger, 6 8 , 1 7 1
leaping. 101
limbs, 1 4 5
locomotion,
Owen, Richard, 19
145
underwater flight, 1 3 7 , 1 3 8 , 1 3 9 , 1 4 0 , 1 9 9
descriptions of plesiosaurs, 1 2 0
dinosaur name, 2 0 , 2 5 , 2 5 n
See
Pinnipeds
also
on ichthyosaurs, 7 1 , 7 2 , 8 7 8 8
Pcntland, Joseph, 6 7 , 1 2 4
Pepys, Samuel, 65
on mosasaurs, 2 3 4
Pbalaradon,
on plesiosaurs, 1 1 9 , 1 2 7 , 136, 1 3 7 , 1 6 0
Phocagroenlandica
85,
8 6 - 8 7
(harp
pliosaur name, 1 6 6
Phospborosaurus,
on pliosaurs, 1 6 9
study of dinosaurs, 2 5 2 6
Pinnipeds
fat,
191,
dawni,
255
Paehyophis
19
247,
problemattcus,
Pacific
Placodonts
237,
turtle
(Dermochelys
coriacea),
41,
140
33,
fossils, 2 0 3 , 2 1 3 , 2 1 9 , 2 2 5 , 2 4 2
lengths, 2 1 4 2 1 5
(Lepidoebelys
olivacea),
34
215
lympanitieus,
Platypterygius,
99,
105,
auslralis,
113
longmani,
113
scientific papers, 6
Plesiopleurodon,
species descriptions, 9 1 1
Plesiosaur fossils
181,
U2(illus.),
113114,
134
192
Palmer, Douglas, 9
Peabody, George, 2 1 2
Pearce, Chaning, 7 1 7 2
183,
jg(illus.),
diets, 2 1 4 2 1 5
249
53(illus.)
180,
3738,
(Plaeodus),
deep dives, 2 3 9 , 2 4 0
248
Peloneusles,
119120
238
210
Plaltcarpus,
245246,
leatherback
236,
Placenticeras,
3536
37(illus.),
3637,
Pistosaurus,
146247
woodwardi,
Pachyrbachis,
Penguins; Seals
See also
<illus.)
53
underwater flight, 1 3 7 , 1 4 0 , 2 3 2
191
Pachycostasaurus,
in
evolution of, 5 7 5 8 , 1 9 9
study of fossils, 1 2 5
Paebyeepbalosaurus,
seals),
214
192
3, 1 3 2
babies, 1 7 4 1 7 5
Pemberton, Ralph, 1 3 0
displays of, 1 2 5 1 2 7
Plesiosaur fossils,
continued
in England, 6 9 , 1 2 0 , 1 2 3 1 2 4 , 1 2 5 1 2 7 , 1 3 2 1 3 3 , 135,
breathing of, 1 2 3
characteristics, 3
in Japan, 1 5 5 1 5 6
descriptions of, 1 1 7 , 1 1 8 , 1 1 9
in Kansas, 1 2 7 , 1 5 7 , 1 5 8 1 5 9
diets, 1 3 4 , 1 5 5 1 5 6
locations, 3
digestive systems, 1 5 6 1 5 7 , 1 5 9
in New Zealand, 1 0 , 1 3 4 1 3 5
earliest, 1 2 0
opalized,
extinctions, 1 4 , 4 4 , 121, 1 6 1 1 6 4 , 2 6 1
i73(illus.)
172175,
eyes, 135, 1 6 0 1 6 1
flippers,
Plesiosaur genera
157158
latest species, 2 5 7
120
Archaeonectrus,
lengths, 1 2 0
i62(illus.)
161,
Attenborosaurus,
limbs, 4 1
130
Callawaysaurus,
132,
Colymbosaurus,
Cryptodidus,
gastralia, 1 4 0 1 4 2
130,
Alzadasaurus,
16,
15
in literature, 2 1 2 2
136
132133,
I33(illus.), I34(illus.),
184
172,
locomotion, 1 3 4 , 1 3 6 1 4 7 , 1 4 8 1 4 9 , 1 5 1 1 5 2 , I58n
175
Dolichorhynchops,
130
Hydraltnosaurus,
130
name, 1 9 , 6 6 , 1 1 8 , 1 2 3 1 2 4
necks, 1 1 8 , 1 2 0 , 1 2 1 1 2 2 , 1 3 5 , 1 5 3 1 5 5 , 1 6 0 , 1 6 8
122
Hydrotherosaurus,
paddles, 1 1 8 1 1 9 , 1 2 0 , 1 3 4 , 1 3 5 , 1 3 8 , 1 4 2 1 4 3
134135
Kaiwbekea,
130,
Libonectes,
136
Kimmerosaurus,
Muraenosaurus,
reproduction, 1 4 7 1 4 8 , 1 4 9 1 5 1 , 1 5 2
159160
ribs, 8 6 , 1 1 9
I22(illus.),
121,
132,
135,
184
130
Polycotylus,
skulls, 4 3 , i33(illus.)
130
Styxosaurus,
skin,161
smallest, 150
Tkalassiodracon,
160161
Tbalassiosaurus,
130
taxonomy, 1 2 9 1 3 0 , 1 3 2 1 3 3 , 1 6 0
Thalassomedon,
23,
129,
Trideidus,
132,
Tuarangisaurus,
Woolungasaurus,
See
also
130132,
i3i(illus.)
135136
10
132,
130
Thalassonomosaurus,
178179
Elasmosaurus;
Plesiosaurus
Plesiosaurs
See also
Plesiosaurus,
124
167
arcuatus,
aquatic adaptation, 1 9 9
bracbypterygius,
conybeari,
161,
184
i62(illus.)
4(illus.\
4,
dolichodeirus,
19,
I2j(illus.),
69,
125124,
133
HAWKMSII,
160
marshi,
diets, 1 4 9 , 1 6 6 , 1 7 0 1 7 1 , 1 7 4 , 1 8 4 , 1 8 7 , 191
eyes, 1 6 9 , 1 7 2
161
2i2(illus.),
198,
houzeaui,
15,
215,
215,
225,
hunting, 1 6 , 1 7 2 , 1 8 0 1 8 1 , 1 9 1
242
latest species, 1 8 9
221
211, 216,
217,
primaevus,
192
tourncnurensis,
Plioplatecarpus,
218,
225,
lengths, 1 7 5 1 7 6 , 1 7 9 , 1 8 5 , 1 8 6 1 8 7 , ' 9 i
232234
limbs, 1 4 2 , 1 6 8 , 183
218
Pliosaur fossils
locomotion, 15, 1 4 4 , 1 4 8 1 4 9 , 1 8 2 1 8 3 , 1 8 9
in England, 1 6 6 - 1 6 8 , 1 6 9 , 1 7 1 - 1 7 2 , 1 8 6 , 1 8 7 , 1 9 1
name, 1 6 6
in Kansas, 1 8 3 1 8 4 , 1 8 8 1 8 9
necks, 183, 1 8 6
reproduction, 150151
in New Zealand, 9 1 0
skulls, 1 7 2 , 1 7 6 , 1 7 9 , 1 8 4 , 193
Pliosaur genera
168,
Archaeonectrus,
i83(illus.),
189,
Dolichorhynchops,
tail fins, 1 6 8
192
180,
Brachauchentus,
190,
183-184,
192,
193
'9
192
time period, 4 4
192
Edgarosaurus,
taxonomy, 1 8 8 1 8 9 , ' 9
Eurycleidus,
192
underwater olfaction, 1 6 9 1 7 1 , 1 7 2
Eurysaurus,
188
vertebrae, 1 7 5 1 7 6 , 1 8 6
132,
Kronosaurus,
I77(illus.),
175179,
180,
185,
192,
weights, 1 7 9 i 8 o n , 1 8 2
See also
'93
172175,
Leptodeidus,
I73(illus.),
Pliosaurus,
180,
'9
169
169,
hrachydeirus,
187188,
192
169,
macromerus,
180,
Mcgalneusaurus,
ig3(illus.)
183,
214
192
214
bennisoni,
181,
Plesiopleurodon,
192
tuckeri,
Plurtdens
192,
Polyptychodon,
Rhomaleosaurus,
169,
187,
188,
i7o(illus.),
171172,
174,
192
Polycotylidac,
189,
192
192
latipinnis,
185
11411%
walken,
193
Polycotylus,
Stretosaurus,
214
192
Polycotylus,
Sitnolcstes,
186
169
rostratus,
Plotosaurus,
180,
Peloneustes,
185,
192
191,
Pachycostasaurus,
185
169171,
hrachyspondylus,
180,
193
major groups, 1 9 1 1 9 2
Maresaurus,
192,
192
andrewsi,
166167,
Macroplata,
192
Pliosaurus
breathing of, 1 6 9
120,
rostratus,
Liopleurodon;
Pliosaurs
166
longirostris,
a/so
Pliosauridae,
5c?
i9o(illus.),
188190,
Trinacromerum,
171
cramptoni,
130
192
184185
192,
Polyptychodon,
definition, 2 7
193
Porbeagle sharks
(Lamna
nasus),
62(illus.),
79
eggs, 2 7
200,
hueni,
Proganockilys
198199,
Prognathodon,
202
31,
quenstedi,
evolution of, 2 9 , 30
32
219,
225,
lack o f stamina, 4 7
229
226227
currii,
ruling, 4 4
227
overtoni,
222(illus.)
221222,
saturator,
solvayi,
ribs, 6 3 n
221
giganteus,
221,
Sphenodontidac,
See also
244
Rbomaleosaurus,
213
Pteranodon,
Pterodactyle,
24,
Pythonomorpha,
208,
2o8n
Rhomaleosauridae, 1 7 2 , 1 7 4 , 1 9 1 1 9 2
255
Protoeeratops,
Rhinosaurus,
6768
Proteosaurus,
8485
terrestrial, 2 9
22611
waiparensis,
ectothermic nature, 2 7 2 8 , 2 9 , 4 5 , 52
zetlandicus,
69
210,
169,
i7o(illus.),
174,
192
171171
Ridley turtles, 34
245
96
Qianichthyosaurus,
Quenstedt, Friedrich, 7 5
227
Rikkisaurus,
Raup, David M 9 , 2 5 8
Riou, Edward, 2 2
Reproduction
Ritchie, Alex, 1 7 3
Robinson, Jane Ann, 1 3 8 1 4 0 , 1 4 1 , 1 4 2 , 1 4 4 , 1 4 6 ,
of aigialosaurs, 2 1 7 2 1 8
o f cetaceans, 3 0 , 7 3
148
o f elasmosaurs, 1 4 7 1 4 8
Rolex, 9 0
o f ichthyosaurs,
43, 4 8 4 9 , 7 2 7 4 , 7 6 , 9 7 9 8 ,
Roosevelt, Theodore, 1
99
Rothschild, B . M., 1 0 9 , 1 1 1 , 2 0 5 , 2 3 9 , 2 4 0
o f mammals, 3 0 , 4 9
of marine reptiles, 1 4 , 2 9 3 0 , 4 8 4 9 , 58
o f mosasaurs, 1 9 9 , 2 1 7 , 2 1 8
of p l e s i o s a u r s , 1 4 7 1 4 8 , 1 4 9 1 5 1 , 1 5 2
Roux, Erica, 2 3 6
Rudwick, Martin J . S., 2 1 , 2 2 , 2 4 , 7 0
Ruling reptiles, 4 4
Russell, A. P., 55
of pliosaurs, 150151
viviparous, 2 9 3 0 , 4 3 , 4 8 4 9 , 5 8 , 7 2 7 4 , 9 7 9 8 ,
99,
217218
Reptiles
anapsids and diapsids, 2 8 2 9 , 5
8
characteristics, 2 7 2 8
Ruta, Marcello, 2 4 9
Ruxton, Graeme D noin
Saint-Hilairc, Geoffroy, 19
See
255
Saltosaurus,
Saltwater crocodiles
(Crocodylus
porosus),
58,
Sander, Martin, 7 6 , 8 2 , 8 7 , 9 2 9 3 , 1 1 4
216
also
Turtles
57
Seilacher, A., 2 3 7 , 2 3 8
Sauropterygians, 3 4 - 3 6
Scanlon, J. D., 2 4 6 2 4 7
Selmasaurus
Schevill, W. E., 1 7 5
Sepkoski, Jack, 2 6 2 2 6 3
Schopf, T h o m a s J. M., 2 6 0
Shannon, Samuel W , 2 1 6 2 1 7
Sharks
217
russelli,
ancient, 2 1 7 , 2 2 1 2 2 2 , 2 4 1 2 4 2 , 2 5 9
Sclerotic rings, 5, 7 8 1 1
attacks on mosasaurs, 2 1 7 , 2 2 1 2 2 2 , 2 4 1 2 4 2
o f ichthyosaurs, 7 7 , 9 6 , 1 0 9
cat-,
of plesiosaurs, 1 6 0
circulatory systems, 4 7 4 8
o f pliosaurs, 1 6 9 , 1 7 2
dogfish, 2 1 7 , 2 1 8
Scotland
7879
ginsu, 2 4 1 2 4 2
great white, 5 9 , 7 9 , 2 1 6 , 2 3 9
horn, 2 2 9
hunting behavior, 2 3 9
Seals
lamnid, 4 7 4 8 , 1 0 3 , 2 4 1 2 4 2
elephant,
111
mackerel, 7 9
harp, m
See
also
mako, 7 9
Pinnipeds
porbeagle, 6 2 ( i l l u s . ) , 7 9
Sea turtles
similarities to ichthyosaurs, 11
tails, 8 , 7 9 , 1 0 2
comparison to plesiosaurs, 3
current species, 3234
88(illus.),
Shastasaurus,
egg laying, 4 7 , 1 4 8
alexanirae,
endangered, 34
altispinus,
extinct species, 31
careyi,
88,
89
89
flatback, 34
earinthiaeus,
green, 33
neoseapularis,
neubigi,
leatherback,
32, 33,
53(illus.),
5354, 58
osnwnti,
pacijieus,
9395
89
89
8889
9293
89
89
Sheldon, Amy, 2 0 5 , 2 4 0
metabolism, 54
Shonisaurus,
ridlcy, 34
gi(illus.)
fossils, 8 9 9 2
Shonisaurus,
continued
Sphcnodontidae, 8 4 8 5
lengths, 4 4 , go, 1 0 4
92
mulleri,
popularis,
gz
Squalicorax
silberlingi,
91
Steneosaurus
tails, g 3
(dogfish
64,
cuniceps,
macrophasma,
quadnscissus,
188,
81,
8$(iIIus.)
82,
75
75
81,
103
Sternberg, George, 2 0 9
Sternberg, Levi, 2 1 3
187
vorax,
74,
Sternberg, Charles, 2 0 6 , 2 1 2 , 2 1 3 , 2 1 8
192
187
nowackianus,
217
66
bollensis,
Stenopterygius,
Simolestes,
shark),
Sternberg Museum. 1 8 4 , 1 8 9 , 2 0 9
Sircnians
Stewart, J. D., 2 0 3
bone densities, 2 4 0
Stinncsbcck, Wolfgang, 9 5 , 1 8 1 1 8 2
Stones
locomotion,
ballast, 1 5 7 , 1 5 9
136
manatees, igi
Slingluff, ] . J., 2 4 g 2 5 0
gastroliths,
Sloanc, Hans, 65
Slowworms
(Anguis
fragilis),
250
174
Storrs, Glenn W , 1 1 8 , 1 2 9 1 3 0 , 1 3 7 , 1 4 9 1
160
Snakes
cctothcrmie nature, 4 5 , 5 2
Strcptosauria,
evolution of, 2 4 5 2 5 0
Stretosaurus
relationship to mosasaurs, 2 4 5 , 2 4 7 2 5 0
Stukcly, William. 19
skulls, 2 4 8 2 4 g
Stupendemysgeographicus,
teeth, 2 4 9
Styxosaurus
See also
Soliar, T.,
2 2 3 . See also
Lingham-Soliar, Thcagarten
127, 210
185.
macromerus,
also
32
130
snowii,
Sues, H.-D, 2 1 4
Sucvolevtatban,
Solnhofen, Germany, 9 , 9 6 g 7 , 2 0 2
Svalbardosaurus,
Sdmmcring, Samuel, i g 6 1 9 7
Sweden
86
80
South Africa, 1 7 4 , 2 2 2
mosasaur fossils, 2 3 0
museums,
South Dakota
See
213
fossil sites, 9
Swinton, W. E., 2 2 2
mosasaur fossils, 1 9 8 , 2 1 6 , 2 1 7 , 2 1 8 , 2 2 8 , 2 4 0
plesiosaur fossils, 1 3 0 , 1 5 7 1 5 8
Spencer, R S., 7 0
Swordfish ichthyosaurs, 1 0 7 1 0 9
l.iopleurod
Taniwbasaurus
227
oweni,
Tbalassonomosaurus
1 8 6 , 1 8 7 . Sec also
130
marsbi,
Thompson, Keith S 1 0 2
Thulborn, R. A., 1 4 9
Thulborn, T , 1 2 0 1 2 1 , 1 7 6 1 7 7 , 1 7 8 1 7 9
Tate, R., 1 6 6
Tokaryk, Tim, 2 1 8
Taylor, M. A., 1 6 0
Tonoglu, C 2 2 0
Triassic era, 11
Tchernov, B 2 4 6 , 2 4 7 2 4 8
Teeth
255
Triceratops,
edcntulousness, 1 1 2 1 1 3
Truleidus
o f ichthyosaurs, 8 7 , 9 8
Trinacrotnerum,
ol
96
Toretocnemus,
Torrcns, Hugh S 6 9
mosasaurs,
107,
198199,
^ 1 s.
223,
224,
226,
228-
229, 234238
189190
bonneri,
188189
willistoni,
Tsujita, C. J., 2 3 7 2 3 8
10
keyesi,
of snakes, 2 4 9
studv
Tucker, W. M., 2 1 4
Tuna
of whales, 1 7 6
to6(tllus.)
Temnodontosaurus,
bluefin, 6 2 / i l l u s . )
circulatory systems, 4 8
106
burgundiae,
192
189
bentonianum,
Tuataras, 2 7 , 8 4
of,
135136
i9o(iIlus.),
Tuarangisaurus
191
132,
seeleyi,
diets, 1 0 5 1 0 6
locomotion,
eyes, 105
tails, 7 9
103
fossils, 6 8 , 81
lengths, 105
Turner, S 1 2 0 1 2 1 , 1 7 6 1 7 7 , 1 7 8 1 7 9
sclerotic rings, 1 0 9
Turner, Thcophilus, 1 2 7
Turtles
current species, 3031
relrapods
emergence from water, 1 3 1 4
limb evolution, 14
shells, jo, 3 2 , 38
Thalasstodracott,
160161
"Bunker," 2 1 6
Tbalassiosaurus,
130
capensis,
Thalassomedon,
23,
banningtoni,
iii(illus.)
129, 130132
222
deep dues, 2 3 9 , 2 4 1
distribution,
215
Tylosaurus,
Wade, M 1 0 5
continued
20711,
dyspelor,
Wahl, W, 1 0 9 , 111
222
fossils, 2 0 3 , 2 0 6 , 2 1 9 , 2 2 0 , 2 4 2 , 2 5 7
Walde, Keary, 8 9
baumuriensis,
z z j
hunting, 235, 2 3 9 2 4 0
Watson, D. M. S., 1 3 7
lengths, 2 1 6
Wei, Chen, 9 6
proriger,
209210
nepaeolicus,
203,
2o6(illus.),
207n,
208, 2o8n,
209(illus.),
189
210, 219, 2 2 8
skulls, 2 0 4
Westermann, G. E. G 2 3 7 2 3 8
teeth, 2 0 3
vertebrae, 2 3 0
207
weights, 2 1 6
zangerli,
Tyrannosaurus,
rex,
Westlothiana,
29
Whales
219
26,
28,
176,
baleen, 5 7 , 93
255
beaching,18
26, 182
Blainville's beaked, 2 4 o n
University of Alabama Museum of Natural History,
blubber, 5253, 5 4
bone densities, 2 4 o n
216217
colors. 8
killer, 5 9 , 101
226
locomotion, 4 1
University of Kansas
Museum of Natural History, 1 5 8 1 5 9 , 203
pilot, 113
primitive, 125
teeth,176
vestigial limbs, 41
78(illus.),
sperm, 53, 5 9 , 1 7 6
7879,
viviparous reproduction, 30
8081
weights, 1 7 9 i 8 o n , i 8 2 n
Whitby, England, 6 8 , 1 2 6 1 2 7
Varanid lizards, 11
W h i t e , T. E., 1 6 6
Varanus
komodoensis
(Komodo
dragon),
58
Whitear, M., 1 0 0
Vega Island, 2 1 5 , 2 1 7
Verne, Jules, 2 1 2 2
W i e n s , J. J., 2 4 9 2 5 0
Vida, Joe, 1 7 2
Wiffen, Joan, 9 1 0 , 2 2 7
Volcanic activity, 2 5 4
Willis. Paul. 1 7 2 - 1 7 ;
188,
198,
205,
206,
211,
213,
216,
217,
239-
Yale University, 1 2 7 , 2 1 2 2 1 3
Yoda, K., 101
You, H.,
240
86
Wimanius,
104
Young, George, 6 8 6 9
Zahcr, Hussam, 2 4 8 , 2 4 9
Wood, Roger, 32
Wcoiungasaurus,
glendowerensis,
178-179
Zalophus
132
Zangerl, Rainer, 2 1 9
Zdansky, Otto, 2 2 2
Zeuglcdon,
Xiphiasgladius
(broadbill
swordfish),
107,
io7(illus.)
232
125