Sea Dragons - Predators of The Pre Historic
Sea Dragons - Predators of The Pre Historic
Sea Dragons - Predators of The Pre Historic
RICHARD ELLIS
© 2OOJ b y R i c h a r d L l l i s
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
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.
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 9 8 7 6 5 4 3 2 1
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 165
T h e Mosasaurs 195
T h e "Reason" for Extinction 253
References 161
Index 291
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 . )
In 1999, because I was working on Aquagenesis, a book about the origin and
evolution of life in the sea, I attended a meeting in Copenhagen enticingly
titled "Secondary Adaptation to Life in Water." I got a lot of material and
references on the evolution of whales, seals, manatees, and penguins, but I also
met N i e l s Bonde and Per Christiansen, participants in the meeting because
they were working on a giant mosasaur skull from Israel that was at the
Geological M u s e u m in Copenhagen (described on page 225 of this b o o k ) . Per
read my description and corrected my misinterpretations. Also in C o p e n -
hagen was Betsy N i c h o l l s of the Royal Tyrrell M u s e u m in Alberta, one of the
world's foremost authorities on marine reptiles. I bothered her in C o p e n -
hagen, and when she thought she had escaped safely to C a n a d a , I found her
there and continued to ask embarrassingly s i m p l e - m i n d e d questions. ( A t the
Copenhagen meeting, Betsy presented an early discussion of the excavation of
the giant ichthyosaur discussed on page 89.)
In 1997, I was invited to Edinburgh to repaint a giant squid model. (I was
then w o r k i n g on a book called The Searchfor the Giant Squid, and I had developed
a preternatural interest in the models found in m u s e u m s around the world.) I
painted the Scottish squid, but more important, I met M i k e Taylor, who is
the m u s e u m s curator of vertebrate paleontology. M i k e was also at the 1999
Copenhagen s y m p o s i u m , and after listening to his presentation, I realized
that I had found the perfect person to alleviate my confusion about almost
everything. I shamelessly badgered him with inane and sophomoric questions,
trying once again to master a subject (the marine reptiles) that I found utterly
fascinating.
One of the first things I did when I c o m m e n c e d this study was the same
thing any student does when beginning a term paper: I searched the Internet.
To my surprise (and, I must admit, satisfaction), almost every quest for
information on plesiosaurs, ichthyosaurs, and especially mosasaurs, produced
the same website: "Oceans of Kansas," the brainchild and production of M i k e
Everhart, now adjunct curator of p a l e o n t o l o g y at the Sternberg M u s e u m of
N a t u r a l H i s t o r y in H a y s , Kansas. M i k e has collected and posted an incred-
ible a m o u n t of information (and illustrations) on the fossils of Kansas, and
although there is much to be found on his website, there was much that I
wanted to know that wasn't there, so M i k e became another victim of my
unfulfillable curiosity. T h a n k s , M i k e , for putting up with me, and congratula-
tions on your forthcoming book called ( o f all t h i n g s ) Oceans of Kansas.
W e ' l l go up the stairs to the fourth floor and pass through the new
Saurischian Dinosaur 1 bill on our way to the Hall ol Vertebrate Origins. It's
tempting to linger here, looking at the a m a z i n g Tyrannosaurus rex or the
duckbilled hadrosaurs, larger than we remember them in the old halls (they're
now standing on raised platforms), but we're on our way to something that I
promise will amaze you. N o w we're in the H a l l of Early M a m m a l s , where we
can see the m a m m o t h s , the mastodons, and the lovely murals by Charles R.
Knight. We pass through an orientation center that has a surprisingly realis-
tic, fully realized little Barosaurus stationed here, a "photo o p " if ever there was
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 resem-
ble 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 astonish-
ing 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
So even though there have been m a n y claims that the Loch Ness monster is a
plesiosaur left over from the past ( a n d that a single animal somehow managed
to remain alive for 100 m i l l i o n years), this is not the skeleton of "Nessie."* It is
* It now appears that the original "Nessie" is a fraud. On March 1 3 , 1 9 9 4 , the London Sunday
Telegraph published this headline: "Revealed: T h e Loch Ness Picture Hoax." T h e front-page
story detailed a complicated plot involving a filmmaker and big-game hunter named
Marmadukc Arundel Wcthcrcll, his son and stepson, and Dr. R. Kenneth Wilson, the man
who allegedly took the famous "surgeon's photograph" that has been the basis for all Nessie
the plcsiosaur Thalassomedon, a representative of a g r o u p of marine reptiles that
thrived for about 100 million years from the Triassic to the Cretaceous
periods and then, for reasons not understood, became extinct. S o m e had long
necks, some had short necks; some were petite, and some were gigantic. S o m e
had sharp little teeth like a python, while others had 8-inch daggers that ri-
valed the fearsome dental equipment of T. rex. Like all reptiles, they breathed
air, but some, unlike any living reptiles, probably gave birth to living young
underwater. (Female sea turtles come ashore to lay their e g g s . ) Sea turtles are
the only living reptiles that have four flippers, so we have to compare the four-
finned plesiosaurs to them, but whereas the turtles arc slow swimmers, some
of the plesiosaurs were aggressive predators and had to chase down their prey.
Turtles are encased in a pair of shells, known as the carapace ( t o p ) and the
plastron (bottom :, and although the plesiosaurs had no shells, they did have a
set of belly ribs called gastralia. H o w they actually swam —whether they "flew,"
"rowed," or performed some combination of the two —has been a subject for
much palcontological speculation. Plcsiosaur fossils have been found all over
the world, including England (the first one was found in 1811 by the famous
fossilist M a r y A n n i n g ) , Kansas, W y o m i n g , C o l o r a d o , Germany, Russia,
Japan, Africa, the M i d d l e East, M a d a g a s c a r , N e w Zealand, and Antarctica. A
plesiosaur fossil found at C o o b e r Pedy in Australia was completely opalized.
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- mingbird is Archilochus colubris, and the box jellyfish is ChironexJlcckcri. T h e first
struction of of these names identifies the genus ( l i o n s and tigers are in the genus Panthera),
Plesiosaurus and the second identifies the species. In all cases, the genus (generic) name is
dolichodeirus,
capitalized, and the species (specific) name is not. Lion, tiger, blue whale, and
one ojthe first
so forth anY the names of these animals in English; in other languages, of
clasmosaurs jound
course, they are different. But whatever the language, the scientific name
in England.
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 Mosa-
saurus 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 long-
extinct 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 in-
stances 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.
In many instances, a single tooth, bone, or bone fragment has been found,
and because there is no other possible explanation, the paleontologist identi-
fies 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, experi-
ence, 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 sea-
side 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 Forma-
tion 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.
T h e marine reptiles all lived in the water. 1 hey all breathed air: some 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 elimi-
nated 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
dolphins is horizontal. T h e mosasaurs were also tail-powered swimmers, but
their propulsion came from sinuous oscillations of the tail, quite different
from the short power stroke of the ichthyosaurs. ( T h e a g a r t e n L i n g h a m -
Soliar believes that at least one mosasaur species — Plioplatecarpus marshi— "flew"
through the water, using its fins as well as its tail, but not many agree with
h i m . ) A n d the plesiosaurs, with their short tails and four powerful flippers,
moved through the water somehow, but the experts disagree as to how this
might have been accomplished. However they propelled themselves, the
short-necked plesiosaurs ( k n o w n collectively as pliosaurs) met n o r m a l re-
sistance from the water; the long-necked ones seemed to have problems that
they were obviously able to solve (a long neck held out in front of a s w i m m i n g
animal would act as a r u d d e r ) , but we still can't figure out how they did it.
Some 65 million years ago, at the geological boundary of the Cretaceous
and Tertiary eras ( k n o w n as the K - T b o u n d a r y ) , a massive asteroid slammed
into the earth at a place that would eventually be identified as Chicxulub, off
the Yucatan peninsula in the Gulf of M e x i c o . Because this impact coincides
with the last recorded terrestrial dinosaurs, there are those who draw a
connection between the two events and claim that the environmental havoc
caused by the impact led to the dinosaurs' extinction. Others hold that
different variables, such as climate change, massive volcanic eruptions, and
elimination of their food source, were at least partly responsible for the
demise of the dinosaurs. N o w it is believed that today's living birds are
actually descended from terrestrial, feathered dinosaurs and that dinosaurs
are not extinct after all.*
* In The Evolution and Extinction of the Dinosaurs (1996), Fastovsky and Weishampel answer the
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 crea-
tures 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."
In a 2002 paper entitled "Extinction of Ichthyosaurs: A Catastrophic or
Evolutionary Paradigm?" L i n g h a m - S o l i a r wrote:
We have reviewed the various stages through which the ancestors of reptiles passed to be
completely freed from the aquatic existence and become purely land-dwelling types. But,
curiously, no sooner did the reptiles attain this terrestrial mode of life than many groups
began to reverse the process and return to the water. We have noted that among the ruling
reptiles the phytosaurs and crocodiles had returned to an amphibious type of existence,
and also among the lizards several groups have become water dwellers.
— Alfred Sherwood Romer
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 exam-
ined 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.
* 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."
In early-nineteenth-century Europe, most educated people believed that all
the creatures God had made had been around since the Creation, so the idea
that there were some that had mysteriously disappeared or died out was
anathema to the prevailing religious doctrines. Georges Leopold Chretien
Frederic Dagobert, Baron Cuvier (1769—1832), was the founder of compara-
tive anatomy and probably the first paleontologist; as the " M a g i c i a n of the
Charnel House," he demonstrated that he could reconstruct an entire animal
from a single fossilized bone. He believed that fossil sequences were the result
of periodic catastrophes, with groups of animals being replaced by new forms
in successive creations, and that each new form was a step in the progressive
sequence that would eventually lead to man, the most sublime of God's
creations. In addition to the biblical flood, he believed that the earth had been
subjected to a succession of natural catastrophes throughout its history.
Cuvier died 30 years before Darwin published his theory of evolution, and
throughout his life, he believed that the extinct types were the ones that had
been swept away by successive disasters, and fossils were the irrefutable evi-
dence of those disasters. In reference to a series of ( i m a g i n e d ) floods, he
wrote:
These repeated [advances] and retreats of the sea have neither been slow
nor gradual; most of the catastrophes which have occasioned them have
been sudden; and this is easily proved, especially with regard to the last of
them, the traces of which are most c o n s p i c u o u s . . . . Life in those times was
often disturbed by these frightful events. Numberless living things were
victims of such catastrophes; some, inhabitants of dry land, were engulfed
in deluges; others, living in the heart of the seas, were left stranded when
the ocean floor was suddenly raised up again; and whole races were de-
stroyed forever, leaving only a few relics which the naturalist can scarcely
recognize.
* 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 wan-
dered 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 pro-
posed periodic mass extinctions caused by catastrophes like the biblical del-
uge, 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 Natu-
rae), 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.
T h e reptilian egg is porous, and the shell contains the chorion, which
surrounds the embryo and yolk sac; the allantois, which is involved in respira-
tion (oxygen in, carbon dioxide o u t ) and stores waste materials; and the
amnion, which lies within the chorion and surrounds the embryo. T h e a m -
nion creates a fluid-filled cavity in which the embryo develops, and the
chorion forms a protective m e m b r a n e around the amnion. T h e allantois is
closely applied against the chorion, where it performs gas exchange and stores
metabolic wastes. T h e presence of these membranes permits respiratory ex-
change between the egg and its environment without desiccation. T h e moist
nature of the membranes is perfect for the developing embryo: too dry, and
the embryo dies; too moist, and it drowns. T h e porous nature of the outer
shell means that it cannot be laid in the water, and even snakes and turtles that
are almost wholly aquatic have to come ashore to lay their eggs. ( A m p h i b i a n s
and egg-laying fishes lay their eggs in the water.) Reptiles differ from the
other major classes of terrestrial vertebrates — birds and m a m m a l s —in that
they lack an internal system for controlling their b o d y temperature and are
largely dependent on the temperature of the environment. Amniotcs include
most of the land-dwelling vertebrates alive today, namely, mammals, turtles,
tuataras, lizards, crocodilians, and birds. Although fundamentally terrestrial,
several types of amniotes, such as ichthyosaurs, plesiosaurs, pinnipeds, sire-
nians, and cetaceans, have returned to the sea.
U n t i l recently, the earliest known amniote was the 8-inch long Hylonomus,
dating from the early Pennsylvanian period (310 million years a g o ) of Joggins,
Nova Scotia, described by J. W i l l i a m Dawson in the mid-nineteenth century.
It is characterized by a lack of openings in the skull behind the eye and was
therefore classified as an anapsid, which simply means "no opening." T h e
primitive nature of Hylonomus m a d e it a candidate for the common ancestor of
later reptiles, but a "supposed amniote" known as Westlothiana was discovered
in S c o t l a n d and has been dated from the early Carboniferous period, 350
million years ago. Although its affinities are in dispute, Benton (1997a) wrote,
"It could perhaps be a reptile, as suggested by some characters at the back of
the skull, but other evidence suggests it might be a microsaur, a g r o u p of
superficially reptile-like amphibians." Believed to be even earlier is Casineria
kiddi, found by an amateur collector in Cheese Bay, Scotland, in 1992 (Caseus is
Latin for "cheese"). Twenty m i l l i o n years later than the primitive tetrapods
Acantbostega and lebtbyostega, it bore certain similarities to them but was consid-
erably smaller and had five digits on each of its limbs instead of seven or eight.
W h e r e a s Acantbostega and lebtbyostega would have been awkward and ungainly on
land — if indeed they ever came ashore — Casineria was clearly designed to
spend more time out of the water. For one thing, it was only about 6 inches
long —not nearly as big or heavy as the others —and its fingers were designed
to flex separately, an adaptation for walking on irregular terrain. Its vertebrae
locked together in such a way as to provide strong support for its body when
it was lifted oft the ground, unlike that of the earlier tetrapods, whose looser
backbones were more like those of fishes (Paton et al. 1999). It is not clear if
Casineria was a reptile, but if it was, it would certainly be listed among the first
of its kind.
Reptiles are divided into two groups, anapsids (turtles and their extinct
relatives) and diapsids. T h e diapsids —reptiles with two holes in the skull
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 rep-
t 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 integu-
ment, 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 oc-
curred 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 evi-
dently 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.
* 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 earliest turtles, known from the late Triassic (200 million years a g o ) , had a propel it through the
shell and an anapsid skull —a solid block of bone, with no openings for jaw water, hut exactly
muscles. T h e first known turtle was Proganochelys quenstedi, which had a fully how it fed with those
' 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 body-
plan 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
T h e steps required to get from a Permian scaly reptile to a fully armored
creature with flippers for feet are also unclear. T h e turtles are probably
descended from reptiles that inhabited swampy areas, where their bodies
widened while their feet became webbed and they lost the articulation of their
digits altogether. (Terrestrial tortoises have clawed toes, not unlike those of
crocodilians.) T h e shell, which consists of an upper element (the carapace)
and a lower one (the plastron), is unique to turtles and tortoises —indeed,
unique in the animal k i n g d o m . T h e path from Permian proto-turtle to recent
turtle or tortoise was not without detours, and there were numerous varia-
tions on the armored reptile theme that d i d not perdure. Proganochelys is no
longer with us, nor are the horned turtles Niolamia from Argentina and
Meiolania from Lord H o w e Island, an isolated speck of land 300 miles east of
Sydney, Australia. In addition to "horns" —actually, flanges of bone behind
the eyes — Meiolania had a spiked tail that was also fully armored. T h e n there
was Stupendemys geographicus, a side-necked turtle that lived only 10,000 years ago
in S o u t h America, whose shell reached a length of 8 feet. In 1976, Roger
W o o d described a specimen from the late Tertiary U r u m a c o Formation of
northern Venezuela that he labeled "the world's largest turtle," but he also
wrote that "Dermocbelys [the leathcrback] is reputedly the largest of all turtles,
living or fossil." It also is not clear if Stupendemys geographicus was fully aquatic.
W o o d wrote, " W h e t h e r it was a fresh water or marine turtle, however, cannot
be determined with certainty from the present evidence. One or perhaps both
pairs of limbs may have been modified into flippers, and the head may not
have been fully retractable."
Today, there are several species of large marine turtles, found mostly in the
tropical and subtropical oceans of the world but occasionally wandering into
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-
replaced by plastic,
the 30-inch-long
hawksbill turtle
(Eretmochelys
imbricata) is still
hunted throughout
its worldwide
its beautifully
mottled shell.
loggerhead turtle
C a i v u . i caret la
has a proportionally
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 turtle (Natator depressus), found only in
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 intcr-
meshed 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 respec-
tive premaxilla immediately lateral to the middle of the rostrum, and as in
Nothosaurus tchernovi, the two anteriormost fangs fit between the two anterior-
most 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.
With Us webbed feet,
Nothosaurus
was a swimmer, but
it was capable oj
tors. Nothosaurs
preceded plesiosaurs
to them.
w a s twice as long as
Hovasaurus plied
the late Penman
characteristics of
both nothosaurs
middle Triassic of Germany, so it seems to have had a limited distribution. and plesiosaurs,
Pistosaurus had a nothosaur-like body with a plesiosaur-likc head, and it has Pistosaurus may
been variously classified as a nothosaur or a plesiosaur, depending on what have been ancestral
38 5 t A WKAgoHS
The placodonts ("plate-teeth") probably used their
something like a
Henodus was
actually an armored T h e term convergent evolution is used to describe unrelated creatures, such as the
placodont. Grooves seagoing turtles and Henodus, that developed similar modifications because
in the jaws may have they lived the same kind of life in the same sort of habitat. Henodus lived
held a baleen-like
during the late Triassic period, and the single species (Henodus chelyops) is
sieve, but how
known only from the Gipskcupcr deposit of Tubingen-Litstnau in southern
Henodus fed—or
Germany.
what it fed on,Jor
a mystery. lizard known as Coniasaurus, from the chalk deposits at Brighton in south-
eastern 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 plesio-
saurs 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 Ichthyosaur was fitted to struggle with the waves of the stormy sea, to
roll therein like modern whales and grampuses, to seize and devour great
fishes, and to dive for them into the depths; and its great, armor-plated
eyes must have been well adapted for vision in the deeper waters. T h e
Plcsiosaur, on the contrary, was fitted for comparatively still and shallow
waters; s w i m m i n g near the surface with its graceful neck curving aloft, it
could dart at the smaller fishes at the surface, or stretch its long neck
downward in search of those near the bottom.
conclusive.
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.
* 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 i°C, 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
In 1964, John Ostrom of Yale University discovered the fossil of a bipedal
dinosaur in the m o u n t a i n s of southern M o n t a n a . From its structure, it was
obvious that Deinonychus had been an agile, swift predator, but because it was a
reptile, and because reptiles were believed to be ( a n d to have been) slow-
moving, sluggish creatures, the only way that this animal could have run the
way it was designed to run was if it had been w a r m - b l o o d e d . Ostrom com-
pared dinosaurs to high-energy m a m m a l s and said, " T h e correlation of high
body temperature, high metabolism, and erect body posture is not accidental.
The evidence indicates that erect posture and locomotion are not possible
without high metabolism and high uniform temperature." One of Ostrom's
students on that M o n t a n a dig was R o b e r t Bakker, who was beginning to
develop the idea of w a r m - b l o o d e d dinosaurs. In a 1968 article in Yale Univer-
sity's Discovery magazine, he wrote, "although much work remains to be done
on dinosaur functional anatomy, the m a m m a l - l i k e posture has convinced me
that these rulers of the M e s o z o i c were fast, agile, energetic creatures that lived
at a high physiological level reached elsewhere among land vertebrates only by
the later, advanced mammals." T h e cause was taken up by Adrian Desmond in
1976 in The Hot-Blooded Dinosaurs and summarized in John Noble Wilford's The
Riddle of the Dinosaur in 1986.
an adult ten-ton dinosaur . . . the rate of temperature change would be very much slower
than a large alligator. Indeed, if the same difference in temperature rise were applied to the
dinosaur (an animal 7 0 0 times greater in body mass than the large alligator) then one might
suppose that it would have taken more than 86 hours to raise the body temperature by 1 C
in the adult extinct giant." Later, Colbert, Cowles, and Bogcrt would reduce their estimates
of the amount of time it would have taken a dinosaur to warm up, but they continued to
believe that large reptiles had to modulate their body temperature by moving in and oui of
the sun.
of growth comparable to that of m o d e r n m a m m a l s . ( T h e role of haversian
canals in bones is not clearly understood, but Bakker wrote, " W h a t e v e r their
role, densely packed Haversian canals are clearly marked 'for w a r m bloods
o n l y . ' " ) Dinosaurs, according to Bakker, "grew m a m m a l fashion; they grew
fast and bred early. And their d y n a m i c approach to quick maturity must have
been one of the most powerful weapons in their adaptive arsenal." ( C h r i s
M c G o w a n disagrees with Bakker's conclusions, saying that Bakker might have
confused haversian bone with fibrolamellar bone, which indicates high body
temperature rather than rapid growth.) And in a study of dinosaur growth
patterns published in 2001, Erickson et al. concluded that "dinosaurs exhib-
ited . . . growth curves similar to those of other vertebrates, but had unique
growth rates with respect to body mass. All dinosaurs grew at accelerated
rates relative to the primitive conditions seen in extant reptiles."
As far as we can tell, the extinct marine reptiles differed from living reptiles
in one way: they lived all their lives in water, a circumstance achieved only by
sea snakes today, f f e m a l e sea turtles come out of the water to lay their eggs;
males hardly ever leave the water.) Of stamina, M c G o w a n (1991a) asks, "why
have reptiles got so little?" and answers by saying, " T h e y have a very limited
capacity for aerobic exercise and are therefore unable to sustain any fast
activities." He does recognize, however, that reptiles can be very fast during
periods of intense activity; "just consider how quickly lizards dart from place
to place, or the speed with which a venomous snake can strike its victim." He
concludes his discussion of stamina with:
S o m e sharks, particularly the lamnids such as the great white, mako, and
porbeagle, have a heat-exchange circulatory system that w a r m s up the m u s -
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.
There are many lizards and snakes, some amphibians, and numerous fish
species that are viviparous, but the e n d o t h e r m i c m a m m a l s are the predomi-
nant vertebrate group that gives birth to living young. ( T h e monotrcmes, an
ancient and anomalous g r o u p of m a m m a l s that includes the platypus and
echidna, lay eggs.) Birds, the other endothermic vertebrates, are all egg-layers,
descended from reptilian ancestors. S i n c e reptiles cannot lay eggs in the
water, and it is clear that most of the ancient marine reptiles were viviparous,
the plesiosaurs, ichthyosaurs, and mosasaurs might have been at least partially
endothermic.
In the 2000 edition of his book History of Life, R i c h a r d Cowen addresses
many of the mysteries of marine reptile metabolism. Q u o t i n g Carrier (1987),
he points out that "many fishes have no problem maintaining high levels of
locomotory performance and exercise metabolism. M a n y sharks s w i m all
their lives without rest, for example. Gill respiration gives all the oxygen
exchange necessary for such exercise levels." But because the early tetrapods
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 simulta-
neously 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 coun-
terparts. "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 hypoth-
esis, 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:
S o m e recently evolved marine reptiles had little trouble making the transi-
tion 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 living
back turtle
(Dcrmochelys
coriacca) can
not the metabolism of whales and d o l p h i n s that enables some of them to live turtles, going as far
as 3,000 feet.
in cold water but rather the thick envelope of blubber that encases them and
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
appearances "the N o r t h Sea, Barents Sea, N e w f o u n d l a n d and Labrador in the
N o r t h Atlantic, and M a r del Plata, Argentina, and South Africa in the South
Atlantic; it also occurs throughout the Indian Ocean, in the Northern Pacific,
to the G u l f of Alaska and south of the Bering Sea, in the southwestern Pacific
to Tasmania and N e w Zealand, and in the southeastern Pacific to Chiloc
( C h i l e ) . " T h e leatherback can also dive to depths of 3,000 feet, where the
water is icy cold. H o w can turtles function under such conditions? T h e y
might be able to thermoregulate like m a m m a l s . Even though their metabolic
rate is low, they can maintain a large temperature differential between their
body core and the surrounding water. As James Spotila wrote (1995): "Deep
dives by leatherbacks appear to be supported by an increased O, carrying
capacity of blood and tissue because the lungs undoubtedly collapse owing to
increased hydrostatic pressure. Hematocrits and hemoglobin and myoglobin
concentration are a m o n g the highest recorded in reptiles, and approach levels
found in diving mammals." Indeed, the increased oxygen carrying capacity,
lung collapse, and hemoglobin and myoglobin concentrations are the same
modifications that enable the deepest-diving m a m m a l s (sperm whales, bot-
tlenose whales, and elephant seals) to dive to such prodigious depths. For
protection from the cold, the cetaceans and seals have a thick blubber layer,
and so does the leatherback. W i t h a large adipose tissue layer and large body
size, leatherbacks "can use changes in blood flow to the skin and periphery to
regulate body temperature such that they maintain w a r m temperatures in the
N o r t h Atlantic and avoid overheating in w a r m tropical waters" ( S p o t i l a et al.
1997). Although we cannot know how (or i f ) the extinct marine reptiles were
able to control their body temperature, in the leatherback we can observe a
living marine reptile and perhaps make some educated guesses about the
ichthyosaurs, plesiosaurs, and mosasaurs.
One clue comes from studies of the energy needs of modern reptiles.
T h e s e show that a s w i m m i n g reptile needs only one-fourth the energy it
would need for walking on land. T h i s difference is due to buoyancy in the
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 .
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:
T h e cetaceans and pinnipeds that evolved long after the great marine
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 devel-
oped 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 ana-
logues 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 leather-
back 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, four-
flippercd ichthyosaurs; there are no 50-foot lioplcurodons or kronosaurs (for
which we can probably be grateful); and there are no huge mosasaurs, crush-
ing 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 sub-
aqueous 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 Creta-
ceous demonstrate an incredible example of convergent evolution, with simi-
lar 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, a
porbeagle shark, a
bottlenose dolphin,
demonstrate similar
evolutionary
solutions to the
problem of moving
swiftly through
ichthyosaur, the
although the
vertebral column in
the ichthyosaur is in
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 slip-
pery 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 ceta-
ceans that were to follow them — and to which they are in no way related — ich-
thyosaurs retained their air-breathing requirements and were therefore re-
stricted 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 discus-
sion of the adaptive process:
M o r e than 245 million years ago the ichthyosaurs terrestrial ancestor m a d e
the plunge back into the watery world. W i t h a body designed for walking
on land, getting around underwater would have been a major challenge. By
shortening and flattening the long bones of its limbs and extending and
covering the existing fingers and toes, evolution carved out a set of paddles
front and rear. Fossils of the earliest ichthyosaurs have five distinct rows of
bones in each. Later, the number varied from three to seven, with up to 100
individual bones in each paddle. But ichthyosaurs needed more than oars
and a streamlined shape if they were to take their place as top predators.
T h e solution was obvious: ditch the reptilian tail and replace it with a
model with a good track record for speed. W h y go back to the drawing
board when sharks had solved that problem millions of years earlier? W i t h
a little tinkering, the ichthyosaurs existing tail could be bent downward
near the end to support the lower fluke of its new tail fin. C o n s t r u c t i n g a
scaffold for the mirror image lobe on top would have been more tricky, but
a blade of cartilaginous material d i d the job. W i t h another l u m p of
cartilage on the back to use as a stabilizing dorsal fin —also similar to a
shark's —the transformation was complete.
that the links hence arising represent real transitions from one branch to another of the
animal kingdom; that through a series of such links . . . that which was once a polypus
becomes successively a mollusca, a fish, a quadruped; an idea so monstrous, and so com-
pletelv .11 variance with ihe structure of the peculiar organs considered in the detail . . . and
no less so with the evident permanency of all animal forms, that nothing less than the
credulity of a material philosophy could have been brought for a single moment to enter-
tain i t - n o t h i n g less than its bigotry to defend it."
* The almost impossible duckbilled platypus had only recently been discovered in Australia
Because they have been extinct for 93 million years, no living ichthyosaurs
were ever found in the "seas and large rivers of our globe," but in the fossilized
skeleton of the ichthyosaur, Young did catch a g l i m m e r of the idea that
D a r w i n would publish 40 years later: that animal species were designed for
their particular function. Predictably, however, he assigned this determination
to God: " S o m e have alleged, in proof of the pre-Adamite theory, that in
tracing the beds upwards, we discern among the inclosed bodies a gradual
progress from the more rude and simple creatures, to the more perfect and
completely organized; as if the C r e a t o r s skill had improved by practice. But
for this strange idea there is no foundation: creatures of the most perfect
organization occur in the lower beds as well as the higher." As S i m o n W i n -
chester wrote, " T h e Reverend Young could not, however, go any further than
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:
* The hook includes poems and dialogue between the workmen who were helping Hawkins
dig, reproduced in dialect. Here is a colloquy between two quarrymen over a plcsiosaur
fossil:
"I wonder what tes."
"O a vicry dragcrn a-maa-bc."
"One that stinged Moses a-maa-bc."
"Here's at'un." A tremendous blow with the mallet.
"How he do zound: I wonder of the stwoonc be holler."
"Tes virc stwoone, vire stwoonc is terrible hard —hit tin agcan Jack."
"There's hes baak-bwoonc."
"An thcr's hes ribs."
situ, as well as drawings made under H a w k i n s ' s direction by landscape painter
John Samuclson Templeton, which p u r p o r t e d to show the various reptiles in
their natural habitat. T h e reconstructions depicted the crocodiliform ich-
thyosaurs s w i m m i n g or hauling themselves out of the water to bask on rocks,
with long, poorly defined flippers and straight tails.
Neither Conybeare nor Dc la Bcche thought that ichthyosaurs came out of
the water—in a drawing caption, De la Beche wrote, "I. communis . . . is
represented on dry land, where it probably never reposed, for the purpose of
exhibiting its form" —but when S i r R i c h a r d Owen included ichthyosaurs in
the outdoor display at S y d e n h a m in 1854, ne
opted for a straight-tailed, rock-
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 ich-
thyosaurians 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.*
I therefore submit that the evidence indicates that these Ichthyosaurs were
viviparous, and were probably produced of different relative bulk in dif-
ferent 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 sug-
gests that this is a distinctive character of certain species.
corpse would almost certainly have caused the death of the mother." T h i s and her babies are
explanation accounts for those fossil ichthyosaurs that appeared to show a conjectural, hut it is
species reached a
length oj about
yjeet.
make sense. T h e r e is a fossil of the ichthyosaur Stenopterygius from H o l z m a d e n ,
Germany, with seven preserved embryos inside the body cavity.*
M a n y ichthyosaurs have been discovered in and around Lyme Regis in
Dorset and W h i t b y in Yorkshire, but the premier site for these fossils is
H o l z m a d e n in Germany. As M c G o w a n (1979a) wrote, " M o r e ichthyosaur
skeletons have been collected from the Lower Jurassic of southern Germany
than any other locality." It was here that the best-preserved fossils were found,
many of them so detailed that the complete outline of the animal was
revealed, including the dorsal fin and the lunate tail. T h e first identification of
ichthyosaur embryos was also m a d e from H o l z m a d e n specimens. W h e n
German zoologist W i l l y Ley (1951) described H o l z m a d e n , he wrote, "There
were numerous small open quarries and a few of them (those that happened
to cut into the epsilon s t r a t u m ) yielded ichthyosaurs. Two hundred of them
per y e a r ! . . . O l d Professor O s k a r Fraas of the M u s e u m of Natural H i s t o r y in
Stuttgart, knew all the details of the 'ichthyosaur business' and wrote a report
about it in 1866," in which he said:
* McGowan ( 1 9 9 1 a ) points out that a baby protruding from the cloaca docs not always
mean that the mother died that way. W h e n a school of false killer whales stranded and died
on a beach in Tasmania, they were buried, but not deeply enough. When the site was
revisited several months later, fetuses were seen partly protruding from the genital slits,
"presumably because of the mounting gas pressure in the carcass."
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 exam-
ined 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, Stenop-
terygius 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."
* 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.
205 m i l l i o n years a g o ) , ichthyosaurs of the Jurassic (205 to 140 million years
a g o ) , and those of the Cretaceous (140 to 85 m i l l i o n years a g o ) . In the first
g r o u p is the yard-long Mixosaurus cornalianus* found mostly in Switzerland
and northern Italy but also in other locations. Other mixosaurs have been
found in Alaska, British C o l u m b i a , Nevada, China, Germany, Turkey, and the
Indonesian island of T i m o r , suggesting that this was one of the most w i d e -
spread of all ichthyosaur genera. In this early form, the vertebral column was
not turned down, so it d i d not have the crescent-shaped caudal fin of the later
species; more likely it had a long, trailing tail fin. Its limbs were modified into
fins, but they had the five-fingered inner structure that showed that they were
descended from land animals, but there is no indication what the ancestors of
ichthyosaurs might have been. In his 2000 discussion of ichthyosaurs, M a r t i n
S a n d e r wrote, "One of the most striking things about the evolution of the
ichthyosaurs is that they appear fully formed in the late Early Triassic without
any known intermediates to terrestrial animals. T h e strong aquatic adapta-
tion of the group w o u l d suggest that it had a long evolutionary history with
ample o p p o r t u n i t y for proto-ichthyosaurs to fossilize. However, the Early
Triassic was marked by rapid evolution rates and the origin of many new
groups as the result of the mass extinctions at the end of the Permian."
* The rules of binomial nomenclature are applied as rigorously to fossil species as they are
to recent ones, and all ichthyosaurs have been given a two-part name, with the generic name
preceding the specific. For convenience, the specific name is omitted in discussions of those
animals when there is only one species. Also, although there are several described species
within a genus such as Ophthalmosaurus (O. icenicus, O. natans, O. canlabrigensis), except in
publications identifying a new species or commenting on the differences within a particular
genus, only the generic name is used. Unless otherwise specified, in this general discussion,
use of a generic term such as Ophthalmosaurus refers to all known species.
The early
ichthyosaurs, such as
the ]-joot-long
developed the vertical, sharklike tail fin a n d had a trailing tail, perhaps with an
Mixosaurus, did
incipient upper lobe. Like m a n y of the ichthyosaurs that followed, Mixosaurus
not have the
had e n o r m o u s eyes, set in a circular framework of bony plates known as the
pronounced upper
sclerotic ring. Primitive crocodilians had sclerotic rings, but m o d e r n crocs and tail lohe oj the later
alligators do not. Living fishes, birds, and some lizards have sclerotic rings, and ones. Fossils oj this
their function is to support the eyeballs. Birds and most reptiles do not have species have been
muscles attached to their lenses and therefore cannot change the focus of their found in China,
eyes. Instead, they compress the eyeball using the sclerotic ring, thus pushing Indonesia, Europe,
Triassic ichthycsaur
Utatsusaurus did
not have the vertical
t a i l J i n that
characterized later
species; it moved by
flexing its entire a c c o m m o d a t e by changing the shape or position of the lens within the eye,
body. It was
but some animals such as snakes a c c o m m o d a t e by squeezing the eye, thus
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.
* 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 advan-
tageous than in the open ocean, where greater speed is required. Later ich-
thyosaurs 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 be-
came 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, par-
ticularly the porbeagle (Laiima nasus) and the great white {Carcbaroion car-
charias). "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, Uta-
tsusaurus 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 swim-
ming 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."
" 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.
this was an immature individual —this specimen bears a striking resemblance
to L. tenuirostris ( C o n y b e a r e ) and was named L. moorei, for its discoverer Chris
M o o r e . T h e incomplete skeleton comprised the skull, forefins, and anterior
trunk, and its size was estimated at about 8 feet. As with most ichthyosaurs of
this type, the eye sockets were e n o r m o u s , and in this specimen, they formed
an almost perfect circle ( M c G o w a n and M i l n c r 1999). In 1998, M. W. Maisch
showed that Leptonectes, Excalibosaurus, and Eurhinosaurus form a cladc for which
the name Leptonectidae was erected, characterized by their long, slender
snouts; elongate, slender fins with three or four primary digits; enormous
orbits; and a tendency to develop an overbite.
Prospecting in southeastern France, H u g o Bucher came across an ich-
osaur fossil in the bottom of a ravine near the village of Le Clapier. T h e
disarticulated skull and some teeth were brought to the M u s c e Geologique at
Lausanne, but Bucher ( a n d M a r t i n S a n d e r ) felt that there was not enough
material for a positive identification. T h e authors finally decided that because
of its size —the skull suggests an animal about 16 feet long —it was probably
Stenopterygius, regarded as the most c o m m o n of the Liassic* ichthyosaurs,
having been found in the U n i t e d Kingdom, Germany, France, Switzerland,
and Portugal (there is a possible record from Argentina, but it is not certain).
We cannot tell whether the fossil record shows that the animals themselves
were widely distributed or whether the fossils have been found because condi-
tions in particular areas are conducive to their appearance. As M c G o w a n
(1979a) wrote, "Attempts to construct geographical ranges for extinct taxa are
always circumscribed by the vagaries of the fossil record, and possibly only
reflect the distribution of good fossiliferous exposures. . . . Ichthyosaurs were
probably highly m o b i l e animals, comparable to present day cetaceans, and I
suspect the complete fossil record would reveal that most taxa were widely
distributed geographically."
* The terms Lias or Liassu arc used in England and Germany for the earliest part of the
lurassic, from about 2 0 5 to 1 8 0 million wars ago. 1 be name probably derives from the
Gaelic word leac, for a flat rock. In terms of standard geology, it extends from the Het-
tangian to the Toarcian and is divisible into the Lower Lias (Hettangian—Lower Pleins-
bachian), Middle Lias (Upper Pleinsbachian), and Upper Lias (Toarcian). The famous
Holzmaden material is all Toarcian; the Lyme Regis material is Lower Lias.
From the preserved
} lolzmaden Shales
T h e identification, classification, and naming of ichthyosaurs is an ongo-
of Germany, we
ing enterprise; as various paleontologists examine and reexamine the m a t e -
know that
rial—which is often fragmentary and sometimes goes missing —the phylogeny
Stenopterygius
of these marine reptiles is stretched, compressed, modified, altered, and cor-
had a large, lunate
rected to such a degree that the subject can only be described as a work in tail and a large
progress, as far from completion as the ichthyosaurs themselves are from dorsalfin. It was
bottlenosc dolphin.
Any attempt to produce a stable taxonomy is circumscribed by the uncer-
tainty of recognizing natural groupings of individuals. It is impossible to
estimate the range of individual variations, and the effects of allomctric
growth ( w h i c h are c o m p o u n d e d by the large size ranges encountered)
m a k e taxonomic conclusions conjectural. Von H u e n e sought a solution to
these problems by extreme taxonomic splitting, but the differences be-
tween his groups were often small, and often within the range which might
be expected for extant species. In contrast to von Huene, I have tended to
l u m p specimens together, rather than referring them to different species.
* 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, allow-
ing 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 quintes-
sential 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 Tu-
bingen, 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 ichthyo-
saurs."* 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 Suevo-
leviatban — 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 Phala-
rodon 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 pal-
a 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 pave-
ment of button-like teeth set in short, massive jawbones. T h e original speci-
men 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 poorly-
known 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 con-
sider 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 J C.H TH r O $ AUK S
1
87
batever its proper
name, Shasta-
Shasta County,
California)
probably looked
pattern, which is
imaginary. It is
looking creature is
Shonisaurus. At
a known length of
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. It was also
unusual in that Us
al. (2001) to conclude that "the ichthyosaur bones were deposited on a deep
enormous front
ocean shelf environment." If they didn't strand, then how to explain the
flippers were the
bodies of so many giant ichthyosaurs in the same place? "One intriguing
same size as the
possibility," say the authors, "is that the ichthyosaurs ate fish or shellfish
rear pair.
tainted with a neurotoxin that paralyzed them." T h e y refer to "mass kills of
modern whales along the coast of N e w England" as a paradigm, but unfortu-
nately for the neurotoxin explanation, these " k i l l s " occurred on the very
shallow beaches that the authors say are absent in the ichthyosaur deaths, and
besides, there is no evidence that these whale kills are in any way related to
shellfish poisoning. (Ichthyosaurs probably ate ammonites, but m o d e r n
whales do not eat shellfish, and although many whales and dolphins cat squid,
these ccphalopods have not been shown to carry poisons toxic to cetaceans.)
C a m p collected the Nevada specimens from 1954 to 1957, with additional
field seasons between 1963 and 1965. He excavated the partial remains of 35
to 40 animals, but not one was complete. He typed up a m o n o g r a p h on
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 accompany-
ing illustrations to his friend Joseph Gregory, which resulted in two publica-
tions in 1976 and 1980 (both appear under Camp's n a m e ) . In the 1980 publi-
cation, 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 ap-
peared 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 con-
tains 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: Shoni-
saurus 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.
ichthyosaur
Cymbospon-
dylus with a
juvenile.
Ichthyosaurs another continent ( L i and You 2002). T h e lower jaw contained teeth only on
delivered their young the forward part, and it has the smallest orbit (and therefore the smallest eyes)
alive, hut it is not of any known ichthyosaur. Cytnhospondylus asiaticus is a new species, younger
known whether they
than the m i d d l e Triassic specimens found earlier.
cared for them. The
Paleontologists do much of their work in the field, but there are occasions
pattern on the tail is
when d i g g i n g through m u s e u m collections also produces some interesting
conjectural, as is the
results. In the collection of the Academy of N a t u r a l Sciences of Philadelphia,
flap at the tip.
Chris M c G o w a n may have found another ichthyosaur that was as big as Shoni-
saurus, 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, ex-
posed as several localities in the N e u q u e n Basin." T h e first Argentine ich-
thyosaurs 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 (Fer-
nandez 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.
tentatively interpreted as "minute scales covering the ichthyosaurian skin, an tion of the ichthyo-
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.
of hearing. . . . It is therefore concluded that the possession of directional
hearing capabilities in the ichthyosaurs is extremely doubtful."
In 2001, with radiographer George Kourlis, paleontologist Ben Kear of the
South Australian M u s e u m performed a computed tomography scan on the
skull of a specimen of the ichthyosaur Platypterygius and found that the tiny
inner ear bones were too thick to detect sound vibrations; therefore, ichthyo-
saurs — or at least the one he examined — m i g h t have been stone deaf. T h e scan
also revealed delicate structures deep in the animal's palate (the pterygoid
bones) that may have been related to the sense of smell, and channels and
grooves that suggest that this ichthyosaur m i g h t have had an electroreccptor
system like that of some fishes and sharks (Perkins 2002). T h e Platypterygius
ichthyosaurs unearthed at H u g h e n d e n in Queensland also revealed a fetus in a
fossilized female, incontrovertible evidence of live birth in ichthyosaurs, as
well as the mother's last meal, which consisted of belemnites, fish, and hatch-
ling turtles.
* 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 unre-
lated (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
that "porpoising" —leaping out of the water when traveling at speed —is an
energy-saving proposition, utilized today by dolphins and penguins. In his
1983 study of the energetics of leaping in dolphins and other aquatic animals,
R. W. Blake wrote, "Leaping is energetically less efficient than s w i m m i n g close
to the surface up to a certain speed . . . after which it is more efficient." (For a
1999 article in the Journal of Experimental Biology, Yoda et al. attached acceler-
ometcrs to Adclic penguins in the S o u t h e r n Ocean and calculated that the
birds expended as much energy leaping from the water as they d i d while
swimming underwater, concluding "that porpoising may be a better strategy
for breathing without reducing s w i m m i n g speed.") Like their swimming,
dolphin leaps arc powered by their tail flukes, but penguins use only their
wings for propelling themselves in and out of the water. T h e downstroke of a
dolphin's tail is obviously capable of propelling a 5 0 0 - p o u n d animal c o m -
pletely out of the water, and even killer whales, which can weigh 10 tons, are
capable of prodigious aerial g y r a t i o n s . T h e side-to-side oscillation of an
ichthyosaurs tail is not an ideal engine for an upward leap out of the water,
but many game fishes such as marhn, swordfish, and even m a k o sharks can
launch themselves into the air, and they too use a vertical tail fin that moves
from side to side. L. B. Halstead (1982) wrote: "In most popular books,
ichthyosaurs are shown leaping out of the water rather like dolphins. T h e
main skeleton was in the lower part of the tail, so the side to side movement of
the tail not only drove the body forward in undulating movements, but also
pulled the tail downwards, which raised the forepart of the body. T h i s meant
that the main propulsive force from the tail would have driven the body
upward towards the surface and might have lifted it out of the water."
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 instrumen-
tal in changing the pitch of the shark's body in the water, and subtle adjust-
ments enable the shark to turn and accelerate quickly. Because the ich-
thyosaurs 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
In 1 9 8 6 , German paleontologist Jurgen Riess published an article in which he argued that
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
In a chapter of the 1994 book Mechanics and Physiology of Animal Swimming,
Judy Massare summarized the literature on ichthyosaur swimming:
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
the end of its tail, it uses it."
could probably not have escaped being inertial h o m e o t h e r m s and they proba-
bly 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 understand-
ing 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 man-
age 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 ich-
thyosaur 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 .
animals we know to be descended from early tetrapods, and of course the foreftipper of
gether so that they form a solid, flipper-shaped mosaic. T h e r e are ichthyo- formed a solid,
saurs with three digits per flipper; a couple with four; many with five; and flipper-shaped
some with six, seven, or eight. T h e foreflippers of Ichthyosaurus breviceps, from 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.
early Jurassic
ichthyosaur
Tcmnodonto-
saurus had eyes
that has ever lived. It several species of Tenuwdontosaurus, the largest of which is T. burgundiae, but it
seems reasonable to was only about two-thirds as large as the gigantic Shonisaurus and had teeth in
assume thai it proportion to its size, suggesting that it fed on large prey items, probably
hunted in reduced including smaller ichthyosaurs.
light conditions. The
O n e of the most c o m m o n ichthyosaurs of the English Lower Lias (Lower
dramatic black and
Jurassic) is Leptonectes tenuirostris, found mostly in Somerset. As suggested by its
while pattern is
name (tenuirostris means "narrow r o s t r u m " ) , this is a long-snouted species, but
purely conjectural.
not as long-snouted as those that would follow. Its jaws were long and narrow
but of equal length, rather like today's franciscana dolphin. T h e southwestern
coast of England has been intensively scoured for fossils since little M a r y
Anning found the first ichthyosaur in 1811, but in Somerset in 1984, an
ichthyosaur was found whose upper jaw extended a considerable distance
beyond the lower, or, as M c G o w a n (1986) described it: "mandible shorter
than skull but exceeding 60% of skull length. Snout extends well beyond
anterior tip of mandible but length of s n o u t . . . not greatly exceeding length
of mandible." M c G o w a n named it Excalibosaurus, both for its swordlike jaw
and for the fact that it was found in the west country, the place of the
emergence of King Arthur's sword. T h e r e seemed to be a tendency for the
upper jaw of Leptonectes to extend beyond the lower ( M c G o w a n 1989c), so one
can postulate a direct evolvement from the geologically older Leptonectes to
Excalibosaurus and finally to Eurhinosaurus, the ichthyosaur that looked much
like a swordfish, only with teeth.
T h e first of the swordfish ichthyosaurs was described by Dr. Gideon
M a n t e l l in 1851, from the U p p e r Liassic of W h i t b y , Yorkshire. He named it
Ichthyosaurus longirostris, but it was later determined that it differed enough from
the other known Ichthyosaurus species to warrant its own genus, so it became
Eurhinosaurus longirostris, the "broad-nosed lizard with a long beak." Eurhino-
saurus has long, slender pectoral fins and a tail that was probably lunate like
that of the broadbill swordfish Xiphias gladius. Today's swordfish has a single,
fleshy pelvic fin, but like all ichthyosaurs, Eurhinosaurus had a pair of hind
flippers. Other significant differences included the absence of gills in the
ichthyosaurs (they were air-breathing reptiles and had to surface to breathe,
whereas fishes breathe w a t e r ) and the presence of teeth in the jaws (the
swordfish is toothless). T h e bauplan of the two is so similar that a layperson
shown silhouettes of Eurhinosaurus and Xiphias would be hard-pressed to dif-
ferentiate one from the other. T h e ichthyosaur was considerably larger than
The 100-miUion-
year-old ichthyosaur
Eurhinosaurus
(top) bears a
remarkable
resemblance to the
modern swordfish
Xiphias gladius.
(Inlike the sword-
had teeth.
the fish, however; swordfish can reach a length of 15 feet ( i n c l u d i n g sword),
but some fossils of Eurhinosaurus longirostris are more than twice that length.
* In their 1 9 6 8 study, "Food and Feeding Habits of the Swordfish," Scott and Tibbo wrote,
"There is a special appeal in studies of food and food-getting among the swordfish and
spearfishes because of the unique spear-like rostral development and its use as a slashing
instrument to maim or injure smaller fishes upon which they feed. The swordfish differs
from the spearfishes (marlins and sailfishes) in that the sword is long and it is dorso-
vcntrally compressed (hence the name broadbill) whereas the spearfishes have a shorter
spear and it is slightly compressed laterally. Thus, the swordfish appears to be more highly
specialized for lateral slashing. Such a specialization would seem to be pointless unless
directed to a vertically oriented prey, or unless the swordfish slashes while vertically ori-
ented, as when ascending or descending." In contrast to almost every other suggestion about
swordfish feeding techniques, Charles O. Mather ( 1 9 7 6 ) wrote, "Essentially a bottom feeder,
a broadbill is believed to use his bill as a tool to obtain crustaceans from their cracks or
attachments and to enjoy crabs and crayfish." In Living Fishes of the World ( 1 9 6 1 ) , ichthyologist
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 ich-
thyosaurs 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 whale—sized 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 pal-
eontologist Frederich von H u e n e . Too bad; Grendelius was a wonderful name.
* T h e giant squid (Arcbiteutbis) is usually said to have the largest eyes in the animal kingdom,
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
The huge eyes of of relative aperture used in camera lenses) for Ophthalmosaurus was the lowest
the late Jurassic of any of the ichthyosaurs (the lower the f-number, the more light the eye
ichthyosaur picks u p ) . T h e y believe that this ichthyosaur had the ability to sec where light
Ophthalmo-
barely penetrated and that it "probably had higher visual sensitivity than a
saurus surest that
cat," an animal justly renowned for its vision in low light. We know that these
it was a night feeder.
big-eyed reptiles were habitual deep divers, because their fossilized skeletons
Only the much
showed evidence of the bends. We have no way of knowing if the squid that
larger Tcmno-
dontosaurus had Ophthalmosaurus fed on were bioluminescent, but because so many living squid
larger eyes, hut it species are, it seems a reasonable assumption, and big eyes with a very low
was three times f-number would be especially useful in picking up flickering flashes of light
the size of from squid in the otherwise pitch-blackness of the depths.
Ophthalmo-
" W h y did some ichthyosaurs have such large eyes?" asked Stuart H u m -
saurus.
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 head-
lights of an automobile . . . with a diameter approaching 25 centimeters ( 1 0 inches) they arc
the largest eyes in the animal kingdom."
suggested, [ b u t ] previous estimates may be even more interesting than they
first appear." H u m p h r i e s and R u x t o n found that harp seals (Phocagroenlandica)
can see at light levels found at approximately 2,000 feet, but because the eyes
of the ichthyosaurs were so much larger than those of harp seals, the ich-
thyosaurs may have been able to see at substantially greater depths. U s i n g the
data of M o t a n i et al. (1999), the investigators found that Ophthalmosaurus had a
sensitivity two and a half to four times that of an elephant seal, so it could
probably see in light that was approximately 25 percent of the m i n i m u m
requirements of the elephant seal. T h e s e large seals are known to forage at
depths of more than 3,280 feet, and H u m p h r i e s and R u x t o n realized that
"visual sensitivity is insufficient to explain why these ichthyosaurs had such
large eyes." But in addition to being able to see at very low light levels, the
large-eyed ichthyosaurs were able to resolve fine detail into an image, indicat-
ing an extraordinarily high level of visual acuity. T h e y concluded: "A further
possible consequence of selection for high visual acuity is the use of visual
signaling or individual recognition between ichthyosaurs, perhaps related to
mating or coordinated f o r a g i n g . . . . In summary, we suggest that the large eyes
of Ophthalmosaurus are the result of s i m u l t a n e o u s pressure for sensitivity, allow-
ing prey detection at considerable depths, c o m b i n e d with pressure for high
acuity, allowing these animals to hunt small, fast-moving prey."
Of all the ichthyosaurs, Ophthalmosaurus was also the most streamlined, with
a teardrop-shaped body tapering to a narrow, pointed snout at one end and a
lunate tail at the other. "Repeated Diving W a s N o t for All Ichthyosaurs" was
the title of an abstract by Rothschild, M o t a n i , and W t h l presented at the
September 1999 Society for Vertebrate Paleontology meeting in Denver. By
examining the bones of post-Tnassic fossil ichthyosaurs, they suggested that
those species with a pronounced downward tailbend ( a n d therefore an effi-
cient heterocercal caudal fin) were probably capable of continuous s w i m m i n g ,
which meant repeated dives. Unfortunately, the researchers found that the
ichthyosaurs that dived the deepest and most frequently, such as Ophthalmo-
saurus, were susceptible to the bends. W h e n animals dive deeply, nitrogen in
the blood is forced into solution by the increased pressure, and as they ascend,
it becomes a gas again, but often as bubbles that remain in the joints. N i t r o -
gen bubbles in the brain can be lethal, but they usually form in other places
THE ] 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 evi-
dence 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 En-
gland, 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
lost their teeth as they matured, or they had perfectly serviceable teeth that
ichthyosaur
were loosely attached (like the teeth of some s h a r k s ) and were lost during the
Platypterygius
shown chasing an
hundreds of millions of years between the time the animal died and the time
ammonite, an
the fossils were discovered. S o m e believe the former explanation, and others
extinct cephalopod. subscribe to the second theory. Ryosukc M o t a n i , who provided this infor-
mation (personal communication 2000), said, "After seeing the specimens
[ K i r t o n ] mentioned, I decided that she was probably right. I have been using
Kirton's reconstruction of the Ophthalmosaurus skull with robust teeth, but I
think 1 belong to the minority at this point."
Teeth or no teeth, these ichthyosaurs were squid eaters, like many d o l p h i n s
and beaked whales today, and they probably sucked them up. Suction feeding
has long been postulated in various living cetaceans, particularly s p e r m
whales and beaked whales ( H e y n i n g and M e a d 1996). In a study published in
April 2000, Alexander W e r t h noted that "several authors . . . have postulated
the use of suction feeding by odontocetes, particularly large teuthophagous
( s q u i d - e a t i n g ) species with blunt rostra and reduced dentition." W e r t h filmed
captive pilot whales that had been stranded and rehabilitated and observed
that "food was often ingested without grasping with teeth." Ophthalmosaurus is
anything but short-snouted, but the beaked whales —whose c o m m o n name
comes from their elongated rostra, which have been likened to birds' beaks —
are largely edentulous, and they are known squid eaters. In most species of
beaked whales, the females and juveniles are toothless, and the males have
teeth that are believed to be used in intraspecific fighting.
By the mid-Cretaceous (105 to 85 m i l l i o n years a g o ) , the day of the ich-
thyosaurs was passing, and only one clearly defined genus remained. As the
sole surviving genus of a long and rich heritage, Platypterygius was one of the
most widely distributed of all ichthyosaurs, appearing in fossil formations in
North and South America, England, Europe, Russia, India, and Australia.
T h e first Australian ichthyosaur remains were discovered in Australia in 1865
near the Flinders River in Queensland. Since then, several other fossils have
been found in Queensland, all of which were described as Ichthyosaurus australis,
but they have since been reassigned to Platypterygius australis and then renamed
again as Platypterygius longmani. T h e s e ichthyosaurs are characterized by their
broad front flippers, and because they have multiple accessory digits, they are
popularly (but incorrectly) known as longipinnate ichthyosaurs. T h e most
complete specimen was found in Queensland in 1934. An 18-foot-long sub-
adult, it lacks all but one tail fin vertebra, the pelvic girdle and hind limbs,
parts of the pectoral girdle, and some ribs. A larger specimen was 23 feet in
total length, probably an adult.
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. They also tried to get along with gizzards instead of stomachs,
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
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 paleon-
tologists 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 pen-
guins? 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 plesio-
saurs 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 struc-
ture 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
skull similar to that of plesiosaurs." T h i s io-foot-long reptile bears some
skeletal similarities to both nothosaurs and plesiosaurs, but, as is nearly
always the case, the skeletal material that would show a transition between the
two groups has not been found. So even though the "transition" cannot be
documented, it seems likely that the pistosaurids are somehow connected
with the evolution of true plesiosaurs and are, according to M c H e n r y (per-
sonal c o m m u n i c a t i o n ) , "highly tempting candidates for the role of plcsiosaur
ancestor."
One of the earliest plesiosaurs (as opposed to one of the earliest found
plcsiosaur fossils) was Archaeonectrus, whose name means "ancient swimmer."
Previously known as Plesiosaurus rostratus ( O w e n 1865), this 12-foot-long species
was found in the early Jurassic formations of C h a r m o u t h , England, and later
in Siberia. Both regions were semitropical 200 million years ago. Archaeonectrus
had a large, elongated head with a narrow snout, only 20 vertebrae in its neck,
and hind flippers that were larger than its forelimbs. W h e n Richard Owen
described Plesiosaurus rostratus in 1865, he noted that some of the centra of the
tail vertebrae were compressed, suggesting the presence of a vertical tail fin.
Plesiosaurs seem to have evolved during the Triassic, some 250 million
years ago, but only began to flourish and proliferate in the Jurassic, 200 to 140
million years ago. T h e earliest forms were probably small, no longer than
about 10 feet from nose to tail, and had comparatively short necks, with about
32 vertebrae. In time, they became larger and their necks became longer, and
the latest forms, from the Cretaceous (140 to 65 million years a g o ) , had
incredible snakelike necks and reached lengths of 47 feet. T h e y had four
paddle-like limbs that look smaller in the Cretaceous forms, indicating a
reduced dependence on maneuverability and an increased emphasis on am-
bush predation. T h e s e marine reptiles have been divided into two major
groups, differentiated by the length of their necks. Traditionally, the plesio-
saurs were long-necked, small-headed creatures, whereas the pliosaurs had
short necks and large heads. A succinct differentiation is given by T h u l b o r n
and Turner (1993):
I he plesiosaurs and the pliosaurs shared the oceans with the ichthyosaurs for
about i^o million years, but the last of these giant marine reptiles died out
around 65 million years ago ( a r o u n d the time of the departure of the d i n o -
saurs), leaving no descendants.
There are, however, some who believe that there is still at least one plesio-
saur s w i m m i n g around in a lake in S c o t l a n d . Although it has been identified
as a dinosaur, a fish, a snake, a sea serpent, a giant eel, an otter, or an elephant,
the Loch Ness monster is most often described as some sort of plesiosaur.
T h e idea of a long-necked, plesiosaur-like creature has caught the fancy of
"Nessie" watchers, and many of the " s i g h t i n g s " describe such a creature,
largely based on the famous "surgeon's p h o t o g r a p h " of 1934 (now revealed as a
hoax involving a sculpted model and a toy s u b m a r i n e ) , which shows an
animal with a long, sinuous neck and a small head. Nessie seems to have
vanished into the mists of hoax and folklore, although there arc many who
travel to Inverness hoping for a sighting. T h e ancient plesiosaurs are all gone,
as are the dinosaurs — unless you believe that birds are their lineal descendants,
in which case there are feathered dinosaurs perched in the trees outside your
windows.
* The 70-vcrtcbrac neck makes Elasmosaurus the longest of the plesiosaurs, but it is not the
longest-necked animal on record. That distinction goes to the terrestrial sauropod dinosaur
Mamenchisaurus, which had a neck that was 46 feet long — equal to the total length of Elasmo-
s a u r u s — h u t Mamcnchisaurus had only 19 cervical vertebrae.
necked elasmosaurs,
Muracnosaurus
gels its name from
muraena, the l a t i n
namefor the moray
Muracnosaurus
was 10 feet long.
In an 1824 letter to De la Beche ( q u o t e d in R u d w i c k 1992) about the newly This is
Plesiosaurus
discovered plesiosaur, Conybeare wrote, " H e probably swam at the surface
dolichodeirus,
and fished with his long neck, or lurked in shoal water hid among the weeds,
described in 1811 by
poking his nose to the surface to breathe and catching all the small fry that
Henry Dc la Beche
came within reach of his long sweep, but he must have kept as much as
and William
possible out of reach of ichthyosauri, a very junior member or w h o m with his Conybeare from the
long powerful jaws could have bit his neck in two without ceremony." skeleton of a "fossil
In 1821, Dc la Beche and Conybeare published "Notice of the Discovery of animal" found by
a N e w Fossil Animal, Forming a Link between the Ichthyosaurus and the Mary Aiming at
"PLESIOSAURUS 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 grandipin-
nins. 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 ver-
tebrate 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 Kan-
sas, 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 accom-
modate 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 pa-
leontologists that would last until C o p e died in 1897.
Did the long-necked In a letter to the New York Herald in 1890, M a r s h described the moment that
plesiosaurs, such as he pointed out Cope's mistake:
Elasmosaurus
platyurus, lie on T h e skeleton itself was arranged in the M u s e u m of the Philadelphia
the bottom, weighted Academy of Sciences, according to this restoration, and when Professor
down with stones C o p e showed it to me and explained its peculiarities I noticed that the
they had swallowed, articulations of the vertebrae were reversed and suggested to him gently
and reach upward?
that he had the whole thing wrong end foremost. H i s indignation was
Or did they hang
great, and he asserted in strong language that he had studied the animal for
head-down to snatch
many months and ought at least to know one end from the other. It seems
their foodjrom the
he did not, for Professor Lcidy in his quiet way took the last vertebra from
surface?
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-
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 his-
tory. 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 Dolicho-
rhynchops 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 compli-
cated, 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 species: Elasmosaurus platyurus, Hydralmosaurus serpenlinus, Li-
bonectes morgani, Styxosaurus snowii, and Thalassomedon banningtoni. Alzadasaurus kan-
sasensis, A. pembertoni, and Thalassonomosaurus marshi form an ontogenetic scries of
Styxosaurus snowii from the S m o k y Hill Chalk, and Sharon Springs M e m b e r of
the Pierre Shale. T h e holotype of the genus of Alzadasaurus is synonymized
with Thalassomedon, leaving Alzadasaurus'eolumbiensis without a generic name, and
therefore a new name is proposed." ( T h a t name is Callawaysaurus, for Jack
Callaway, a vertebrate paleontologist who died in 1997.)
impossible to picture
Thalassomedon
moving through the
water. The
a rudder on the
as it paddled through
constant course
corrections.
seen in the Denver M u s e u m of N a t u r a l History, and because the Denver
specimen has been licensed to Valley Anatomical Preparations of Chats-
worth, California, casts of the C o l o r a d o elasmosaur have been sold to the
Royal Tyrrell M u s e u m in Drumheller, Alberta; the M i l w a u k e e Public M u -
seum; and, in Japan, the Iwaki M u n i c i p a l M u s e u m , the Toyohashi M u n i c i p a l
M u s e u m , the M i e Prefecture Natural H i s t o r y M u s e u m , and the Nagoya C i t y
A q u a r i u m . T h e cast in the H a l l of Vertebrate Origins in the American
M u s e u m of N a t u r a l History, discussed in the introduction, was also made by
Valley Anatomical Preparations.
Another spectacular elasmosaur is the Australian Woolungasaurus glendoweren-
sis, n a m e d for W o o l u n g a , a reptile in Aboriginal mythology, and the Glen-
dower 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 Plesio-
saurus 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. richard-
50/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 speci-
men, 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
Reconstruction oj the skull of the plesiosaur Cryptoclidus eurymerus
SA = surangular; SQ = squamosal.
SA = surangular; SQ = squamosal.
arrangement of
powerful muscles of the paddles, which sometimes contained as many as 18
Cryptoclidus
suggests that this 11-
joints per digit. ( O n l y some ichthyosaurs had more phalanges; Platyptcrygius
foot-long plesiosaur had as many as 30.) As reconstructed in Brown's paper, Cryptoclidus eurymerus
fed on shrimps and was a heavy-bodied, long-necked plesiosaur with a shortish tail and a broad,
other small prey lightly built skull, with as many as 100 small sharp teeth that interlocked
items by trapping outside the jaws. T h i s u n c o m m o n arrangement is believed to have functioned
them in the
as a sort of trap for fishes and soft-bodied ccphalopods. Brown showed it
intcrmeshed teeth.
"flying" underwater, with the forelimbs raised on the upstroke and the hind
flippers trailing behind: "the flattened body shape provides additional dorso-
ventral 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 Kai-
wbekea 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, Elas-
mosauridae, 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 ver-
tebrae "provide no evidence of a serpentine mobility, although the craniocer-
vical 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.
* Robinson's paper, published in the Neues Jahrbuchjur Geologic una Palaeontologie in Stuttgart, is
1
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.
other subaqueous fliers ( o t a r i i d pinnipeds, penguins, and sea turtles) to
plesiosaurs and then, after much structural, osteological, and muscular anal-
ysis, concluded that all four limbs of plesiosaurs functioned as propulsive
hydrofoils; that plesiosaurs were probably unable to emerge on land; and that
forces distributed through the body include those that arise from the restrain-
ing actions of major l i m b muscles arising from the inner and outer surfaces of
the girdles, and are absorbed directly by the ventrally located platelike girdle
elements. She concluded that other propulsive forces result from the hydro-
d y n a m i c thrust that tends to drive the limbs forward and are transmitted by
the "bowstring" construction of the ventral basket. " T h e archer's bow con-
struction or the dorsal and ventral elements of plesiosaurs is able to accom-
m o d a t e a s y m m e t r y of force direction and m a g n i t u d e in anterior and pos-
terior, as well as bilateral, wing pairs without i n t e r m o b i l i t y of bony elements
or distortion of the body as a whole."*
Gastralia, also called gastric or belly ribs, have been defined as ossifications
in the belly wall. T h e y lie just posterior to the s t e r n u m and provide sites for
muscle attachment as well as support for the abdomen. In living animals, they
are found in some lizards, crocodiles, and the tuataras, but whereas the
theropod dinosaurs had them, the ornithischians and most of the sauropods
did not. ( M o d e r n birds lack gastralia, but Archaeopteryx and other primitive
birds had a full set.) It is likely that muscles linked to the gastralia were used
to expand the volume of the body cavity in breathing. Gastralia were present
in m a n y of the earlier marine reptiles, such as placodonts and nothosaurs, and
although they were retained by later forms, particularly the ichthyosaurs and
plesiosaurs, they were absent in mosasaurs. In most Jurassic ichthyosaurs, the
gastralia were fairly small, but they were larger in some of the Triassic forms
and much larger in the plesiosaurs. W h e n it was thought that plesiosaurs
might have occasionally emerged from the water — perhaps to lay eggs — it was
suggested that the gastralia provided support for the body on land, but now
" McGowan ( 1 9 9 1 a ) wrote that "the relative narrowness of an inclined plane is expressed as
the aspect ratio which is the ratio of the length to the width. A plane 10 units long and 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.
verable goshawk, have a comparatively low A R . T h e plesiosaurs, with their
h i g h - A R flippers, "arc inferred to have hunted by cruising long distances
searching lor small, dispersed food items, [ w h i l e ] pliosauromorphs are in-
ferred to have actively hunted single, large food items on the wing" (O'Keefe
2002c). But whereas the aspect ratios of fighter planes and long-range b o m b -
ers are specifically designed to achieve their particular purposes, no such "de-
sign" was responsible in the various plesiosaurs, which developed along dif-
ferent lines and achieved different results. T h e s e creatures d i d not develop
their particular aspect ratios so that they could specialize in a certain type of
prey capture; rather, the flipper types evolved over time, and the animals
developed attack strategies commensurate with their capabilities.
In 1989, Beverly Halstead published an article simply titled "Plesiosaur
Locomotion" in which he proposed that the plesiosaurs used all four fins for
propulsion, and because theirs was an undulatory trajectory through the wa-
ter, their motion was not affected by the turbulence created by the vortices of
the front flippers' power stroke. He wrote: "Once an undulating or porpoising
mode is acknowledged then plesiosaurs can be allowed to swim like sea lions."
T h i s is "Newman's porpoising solution," which was in fact proposed by N e w -
man and Halstead himself in a popular article published in 1967.* In this
article (which is nominally about the discovery of a Liopleurodon skeleton in
Bedfordshire), the word porpoising does not appear, but the authors wrote:
* To the eternal contusion of biographers and bibliographers, the man born Lambert
Beverly Halstead : 1 9 ^ — 1 9 9 1 ! was .11 one point given the name of his stepfather and became
Lambert Beverly Tarlo. In palcontological publications from 1 9 5 4 to about 1 9 6 8 , he used the
name Tarlo. Then for a brief period he used both names and became Halstead Tarlo, but
from 1 9 6 9 onward, he used only Halstead (Sarjeant 1 9 9 } ) . This change led to some rather
peculiar constructions, such as this one from "Plesiosaur Locomotion" (whose author is
L. B. Halstead): "On the basis of an unusual bone identified as a scapula. Tarlo ( 1 9 5 8 )
reconstructed the musculature of a giant plesiosaur." This was obviously intended to
identify the reference, but it presupposes that the reader will know that Tarlo and Halstead
arc the same person, which might not be evident to those unfamiliar with Halstead's
convoluted nomcnelatural history.
conclude that these creatures were unable to dive after their prey. Further-
more, we can see from the structure of the shoulder girdle that the associ-
ated 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 pliosaur—it did not have sufficient muscle power for an
effective backing stroke.
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 move-
m 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 acceler-
ating 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
strokes. M o d e r n penguins are equipped with only one set of propellers —
those of the forelimbs ( w i n g s ) —and yet penguins are exceptionally fast, quick
in turns, and capable of dramatic bursts of acceleration. Plesiosaurs, endowed
with two sets of propellers, fore and aft, must have exceeded the penguins in
the top speed possible for a given body size in the capacity for maneuvers in
three dimensions." Of course, not every activity required high-speed s w i m -
ming, and the animals might have been able to change their "gaits," similar to
the way a horse varies its leg movements when walking, trotting, or galloping.
Also, the requirements of the long-necked plesiosaurs were probably quite
different from those of the pliosaurs; the former were mostly fish eaters that
plucked their prey from schools, whereas the latter were large-prey predators
that had to chase their prey down and thus needed more maneuverability as
well as more speed —especially if the prey was as mancuverable and fast as
they were. As N e w m a n and Tarlo (1967) wrote, "From studying the muscle
actions in the two kinds of plesiosaur we get a picture of two different ways of
swimming that we never would have suspected from the general appearance of
the skeletons alone. T h e long-necked forms were adept at twisting and
turning with great speed, but all their time was spent at the surface of the sea.
T h e pliosaurs were powerful swimmers that hunted down their prey; al-
though they did not have the agility of their cousins, they were adapted to
diving down after their food."
" In response to niv question about the word atheoretical i n his title, Lingham-Soliar answered
as follows: "I use atheoretical essentially to mean lacking sound biological or functional
theory. It was not, however to impugn or undermine previous discussions on four-wing
flight but simply to state that despite many of the ideas proposed being good, they
frequently lacked theoretical basis (i.e. they were more empirical, often without due regard
for system and theory). By this I mean anatomical and morphological theory (musculature
and joint construction) was often ignored when they didn't coincide with four-wing flight
hypothesis, body weight and lifting surface areas, despite all the postulations, were never
investigated, anterior and posterior wing shape and design were taken for grained. Sight
theory in living animals and machines was poorly addressed, the bowstring hypothesis was
Recent workers have regarded the four wings of plesiosaurs as four organs
performing essentially the same function, i.e., an integration of the com-
ponents of thrust, lift, stability, and steering in one and the same system. In
extant vertebrates on the other hand (e.g., sharks, whales, dolphins, ich-
thyosaurs, 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 indepen-
dent 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 num-
ber 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.
Traditionally all four plesiosaur limbs have been treated as identical struc-
tures. 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 lower-
ing 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, necessi-
tated by mechanical and anatomical limitations, but every bit as unique.
Superbly suited for life at or near the surface, elasmosaurs feed in one of
two ways. R i s i n g quickly from the depths, snaking their heads through the
water, they can shoot their heads up to twenty feet out into a school of
small fish or squid. Or, by lifting their heads high above the surface, they
can drop down onto their unsuspecting prey. Long, needle-sharp teeth
pierce their victims and seldom allow an escape. W i t h a quick jerk of the
head, the reptile flips its slippery prey up to dislodge it from the sharp
teeth, then catches it in mid-air. A few adjustments, and the fish is poised
to go down the long neck headfirst, so the fins and scales can't injure the
lining of the e l a s m o s a u r s throat.
Next to the little pliosaur fossil was a gaping hole that M a r t i n described as
"extensive evidence of vandalism," suggesting that the site had already been
vandalized and that perhaps some indications of an adult had been removed.
Poaching of fossiliferous sites is bad enough, but in this case, the only
conclusive evidence for live birth in plesiosaurs might have been removed
before a scientist ever got to see it.
Rumors abound within the paleontological c o m m u n i t y about undescribed
plesiosaur embryos in unopened crates, but so far, none of these mysterious
crates has been opened, and the contents —if they exist —remain hidden from
sight and from science.*
Plesiosaurs have raised the art of paleontological guesswork to new
heights. We do not know how these great reptiles hunted, how they swam, or
how they gave birth. Even with sufficient fossil material, we are unable to tell
how the plesiosaurs actually used their four flippers for propulsion. ( T h e
ichthyosaurs also had four flippers, but their s w i m m i n g was powered by their
" 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 arrange-
ment in their beaks. Cormorants and anhingas (the latter sometimes known as "snake-
birds" 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 serpen-
tine 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 plesio-
saurs, 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 anoma-
lous 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
mosasaurs lurking around, the long-ncckcd elasmosaurs must have lived in
constant danger of decapitation.* C o u l d they, as Halstead suggests, "escape
the attention of predators" by s w i m m i n g with the head and neck lifted high
out of the water?
M a n y scientists do not believe that a long-necked plcsiosaur could actually
lift its head and neck out of the water. In his popular article on plesiosaurs,
M i k e Everhart (2000b) wrote:
Unless the laws of physics were suspended on the behalf of these extinct
creatures, it would have been impossible for them to lift much more than
their head above water to breathe. If you would like to prove this for
yourself, try lifting something long and heavy from one end while floating
in water and not touching the bottom. As you try to raise the object, your
feet and the lower part of your body will also begin to surface to counter-
balance any weight above the water. In elasmosaurs, the weight of the long
neck (as much as a ton or more in an adult a n i m a l ) dictated that the center
of gravity was just behind the front flippers. H o l d i n g the head and neck
above the surface was something that could never happen unless the rest of
the body was setting on a firm b o t t o m in shallow water (a possibly fatal
situation for the clasmosaur.'). Even then, the limited movement between
the vertebrae, limited musculature, and the sheer weight of the neck would
severely limit the height to which the head could have been raised.
4
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.
River in H o k k a i d o , consisted of the right h a l f of the pectoral girdle, a nearly
complete right forelimb, disarticulated vertebrae, several gastralia, and frag-
ments of the pelvic girdle. T h e skull was nowhere to be found, making a
specific identification impossible, but the skeleton indicated an animal ap-
proximately 10 feet long ( S a t o and Tanabe 1998). Like living squid and
nautiluses, ammonites had beaklike jaws, and in the region that corresponded
to the plesiosaurs gut, researchers found about 30 a m m o n i t e jaws, suggesting
that the tiny jaws ( n o n e larger than Vi i n c h ) were the contents of the
plesiosaurs stomach. W i t h their slender teeth, unsuitable for crushing the
sturdy shells of ammonites, plesiosaurs had to have swallowed the ccphalo-
pods whole, but curiously, there was no sign of the outer shells.
In Samuel Wilhston's 1903 discussion of N o r t h American plesiosaurs, he
mentioned that "pebbles" were often associated with plesiosaur remains.
" W h a t the use of these pebbles was I will not venture to say," he wrote, "they
may have served as a sort of weight to regulate the specific gravity of the
animals or they may have been swallowed accidentally." U p o n reading W i l -
liston's report, dinosaur collector Barnum Brown (1904) wrote a note for
Science in which he said, " T h e conclusion seems evident that invertebrate
animals formed a large part of the food of plesiosaurs and that, in default of
crushing teeth, the breaking up of the food was effected by the aid of these
stomach stones, the presence of which further implies a thick-walled, gizzard-
like arrangement in the alimentary canal." Harvard University paleontologist
Charles Eastman (1904) was so offended by Brown's remarks that he submit-
ted a rebuttal to Science, writing that "the history of the gizzard shows that it
was developed first amongst cold-blooded vertebrates, then lost by them and
afterwards acquired independently by birds." He concluded: "For our part, we
are w i l l i n g to consign to birds the exclusive enjoyment of gizzards and feath-
ers. . . . Before asking us to believe that all plesiosaurs had 'gizzard-like
arrangements' let it be shown that all plesiosaurs and related reptiles had the
habit of gorging themselves with foreign matter to the extent asserted of the
American species, and let no doubt remain that these pebbles are not of
adventitious origin." T h e last word in this 1904 donnybrook went to the man
who had started the whole thing, Samuel W i l l i s t o n (1904b). He too wrote a
letter to Science in which he said, "Apropos of Dr. Eastman's letter . . . permit
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
* These authors also had no doubt about the way plesiosaurs locomoted: "His paddles are
highly specialized oars. The great development of the head of the prcpodials indicates a
considerable use of these powerful propelling mechanisms. 7 hey were undoubtedly 'feath-
ered' on the return stroke, and then turned with the leading edge down for the propulsive
stroke. It has been shown that the animals had a flexible streamlined body and that normal
locomotion was probably undulatory, the paddles used probably as starters and for sudden
accelerations. The relatively short, stout tail undoubtedly was a powerful organ of propul-
sion." This seems rather unlikely.
M u s e u m at the University of Kansas) and M i k e Everhart, and arrangements
were m a d e to perform an exploratory dig at the site. T h e plesiosaur, cataloged
as K U V P 129774, was not identifiable from the bones, but it was clear that
the stones were gastroliths. T h e y were among the largest ever found, and,
according to Cicimurri and Everhart (2001), they were "even larger than those
reported Irom the giant sauropod Scismosaurus." ( I he largest ol the stones was
more than 6 inches long and weighed 3 'A pounds.) M i c h a e l Taylor's idea that
the "gastroliths can be eaten or vomited to change buoyancy q u i c k l y " d i d not
convince Cicimurri and Everhart, because "it is doubtful . . . that this would
have been a useful strategy for an clasmosaur with a 5—6 m (16—20 ft.) neck,
especially one that was living and feeding hundreds of kilometers from the
nearest sources of such stones." As to the function of the gizzard stones,
Cicimurri and Everhart concluded: " T h e location of the gastroliths within
the abdominal region . . . and their intimate association with the remains of
small fish supports the hypothesis that these stones aided the plesiosaurs in
the breakdown of food."
* The British Museum was created by an act of Parliament in 1 7 5 $ , and its Natural History
Department was formed from collections bequeathed by Sir Hans Sloanc. In 1 8 8 1 , when Sir
Richard Owen superintended the building of the new structure in South Kensington, it was
known as the British Museum (Natural History). In 1 9 6 3 it officially became the Natural
History Museum, London, or, more popularly, the N H M .
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 collec-
tion 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."
century description
of a plesiosaur was
turtle." This is
Plesiosaurus
conybeari, whose
(98.9 to 93.5 million years a g o ) , the last of the ichthyosaurs (the Platy-
p t e r y g i d a e ) were gone. Sixty-five million years ago (the end of the Creta-
ceous), 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 ich-
thyosaurs could explain vvhv thev have not been directly affected," but what-
ever 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 plesio-
saurs 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 bound-
ary—and 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.
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. The capacious chest and great ribs show a
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 sea-
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 Mac-
roplata ("long blades," named for its scapulae), which was either a short-necked
plesiosaur or a (relatively) long-necked pliosaur. Based on its powerful fore-
limbs (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 means simply "scalizard" (from the Greek enalios, meaning "of or from the sea")
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
a tryst. There is
nothing in thejossil
record to indicate
had a shoulder girdle with coracoids that were proportionately much larger species, hut it was
than the scapulae, indicating a powerful forward stroke for fast swimming. certainly possible.
As Darren Naish wrote in a letter to me, " M o s t workers now agree that there Both sexes were
is no simple dichotomy between 'long-necked' and 'short-necked' forms, about ij feet long.
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 plesio-
sauroids . . . [which is] important and interesting in itself, as further study of
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 Eu-
rycleidus 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 short-
necked plesiosaurs of N o r t h America, Kenneth Carpenter wrote: "Tradi-
tionally, all short-necked plesiosaurs have been grouped together into the
Pliosauridae. However . . . the term 'pliosaur' should not be used indis-
criminately 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 aban-
doned to avoid any phyletic implications."
" The Blue Circle Cement Works has proven to be a rich source of vertebrate fossils. A list
reproduced in Grange ct al. ( 1 9 9 6 ) includes fragments of two turtles, several ichthyosaurs,
some dinosaur parts, and, in addition to teeth and bones from other phosaurs (including
l.ioplcurodon), a complete skull of P. brachyspondylus in 1980 and another almost complete
skeleton of the same species in 1 9 9 4 .
From an analysis of
gative paleontologists food, including fishes, cephalopods, and other reptiles as available. It was
have deduced thai the apparently a d o m i n a n t carnivore which despatched large prey such as fishes
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
hunted by smell, the
posterior dentition to help move the prey down the gullet." In other words,
way most sharks do
P. brachyspondylus was a formidable predator, capable of attacking and eating
today. It is thought
anything and everything that it encountered —including dead dinosaurs. Dur-
that other plesiosaurs
ing the excavation, three armored scutes (horn-covered bony plates set in the
relied primarily on
sight.
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:
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 thou-
sands of pieces, was found by a miner named Joe V i d a , whose clumsy at-
tempts 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," the
called "Eric the Half-a-Bcc." ( T h e opalizcd fish gastroliths and vertebrae that plesiosaur with an
opalized skeleton,
were found in the gut regions were n a m e d "Wanda.") Entrepreneur S i d Lon-
once looked
dish bought "Eric" and paid W i l l i s to restore it, but he went bankrupt and
something like this.
put the opalizcd fossil up for public auction. Because the entire fossil was
Leptocleidus was
made of precious opal, its value was estimated at $300,000, and there was no
about tofeet long.
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
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 spon-
sored 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 cephalo-
pods 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:
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 1931—1932, 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 Kro-
nosaurus 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.*
* 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 the
early Cretaceous of
specimen has been restored in plaster." Plaster or no plaster, it is a curious
Australia, was one
thing that "we still do not have a description of the skeleton, despite its
oj the most fearsome
relative completeness" ( L o n g 1998).
predators that ever
Colin M c H c n r y of Canowindra, N e w South Wales, has a thing about
lived on Earth. Its
giant pliosaurs and wrote his Ph.D. dissertation on Kronosaurus. In a letter to massive, jlat-lopped
me in 2002, he said: skull was 9 jeet
with big flippers, Kronosaurus was more of a sea lion with the skull of a croc. carnivores.
teeth of a crocodile
foot-long
Liopleurodon
was the terror oj the
Europe. 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 impli-
cated 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 elas-
mosaur 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 preda-
tor with exceptionally large teeth, probably Kronosaurus. Of the possible en-
counter, they wrote:
* Following the airing of the television series and the publication of the accompanying book
(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
In any event, the description of Liopleurodon's "crown of dagger-like teeth"
appears to be incorrect, since N o e (2001) has shown that this "spatulatc
rosette" of teeth did not exist in life and was caused by crushing of the fossil.
O t h e r large pliosaurs, such as the recently described Maresaurus coccai from
Argentina (Gasparini 1997), were said to have had a spatulate rostrum fitted
with a rosette of large, caniniform teeth, but there too, the splayed-out teeth
did not exist in the living animal. Maresaurus, which means "sea lizard," reached
a length of 20 feet. W i t h its mouthful of sharp teeth and its powerful paddles,
these pliosaurs probably resembled t o d a y s great white shark in their attack
strategics; they rushed at their victims from below and took great bites out of
them. N o e (2001) uses the term "bolt-shake feeding" to describe the tech-
nique of pliosaurs that took huge chunks out of their prey by violently
shaking their heads while holding on with their teeth. W h e t h e r they were 30,
40, or 50 feet long, there is no question that the large pliosaurs—Liopleurodon,
Kronosaurus, Maresaurus, Bracbaucbenius, Megalneusaurus, Pliosaurus, and Peloneusles —
with their massive jaws filled with big, sharp teeth and their short, heavily
muscled necks, were probably close relatives and the predominant marine
predators of their time. Here's how Haines described an imaginary encounter
between 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 oj the giant pliosaur
Liopleurodon
adjusts her limp corpse in his mouth, repeatedly biting and shaking it.
(after Newman and
Eventually it breaks into three pieces and, grasping the front portion, he
Tarlo 196y). Notice
rises to the surface, flicking to the back of his gaping pink throat and
the prominent belly
swallowing.
ribs (gastralia).
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 Liopleuro-
don"), 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 vil-
lage 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 -
shop on Vertebrate Paleontology, held in Florence in September 2001. W h e n
the press got hold of the story, the vertebrae (about which Frcy ct al. had said
"a precise identification is not possible for the m o m e n t " ) had somehow
grown into a nearly complete 65-foot-long skeleton of Liopleurodon, possibly
the biggest marine reptile that ever lived, with "machete-sized teeth and jaws
powerful enough to chew through granite" ( B B C N e w s , December 30, 2002).
A granite-chewing pliosaur may be a bit over the top, but a M e x i c a n pliosaur
(even if it is not Liopleurodon) is indeed newsworthy, as no comparable fossils
have been described from there. M u c h more work is required on the fossil
known as the " M o n s t e r of Arambcrri" ( h a l f of it is still in the g r o u n d ) , but it
is possible that in life this giant pliosaur weighed as much as 50 tons, about
the weight of a full-grown bull sperm whale.* Evidence of a massive bite mark
on the skull suggests a battle between this monster and another, and if this
one was the loser, imagine how big the winner must have been. It appears that
the great marine reptiles are on the way to achieving a reputation previously
accorded only such inftmous terrestrial carnivores as Tyrannosaurus roc—but
no T. rex weighed more than 7.5 tons.
* Animals supported by water and therefore spared the debilitating pull of gravity can
achieve greater weight and bulk than their terrestrial counterparts. T he largest and heaviest
land mammal alive today is the African elephant, which can reach a weight of 8 tons, but the
sauropod dinosaurs were the heaviest animals ever to walk the earth; Colbert ( 1 9 6 2 )
estimated the weight of Braekiosaurus at 78 tons. (It was originally believed that such gigantic
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-foot-
long 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 one of the last of the
buoyant m e d i u m , and an enlarged head m o u n t e d on a shortened neck allowed great mega-predators
large prey to be t a c k l e d . . . . T h e large head, with massive jaw muscles to exert of the late
Cretaceous.
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, hard-
boned 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
paleontology." T h e short neck and relatively long skull are pliosaurian fea-
tures, 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 Cenomanian-
Turonian (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 short-
necked 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 plesio-
sauroids, 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 fre-
quently broken and worn, "suggesting a diet of fleshy prey with fairly large
bones, such as very large fish and other reptiles."
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 . )
T h e latest salvos were fired by Darren Naish, Leslie Noe, and Dave M a r t i l l ,
who wrote an article for Dino Press, a Japanese dinosaur magazine. As trans-
lated into English, the article reintroduces the giant pliosaurs that reached
stupendous sizes. A massive lower jaw was found in the Kimmcridge Clay
Formation and assigned by Beverly Tarlo (1959a) to Pliosaurus macromerus.
Extrapolating from the mandible, the skull would measure "an impressive
2.9 m [9.5 feet]. C o m p l e t e pliosaur skeletons suggest that the total length of
these animals is about six times longer than the total length of the skull. It
appears possible, therefore, that the Berkshire mandible represents an animal
of an a m a z i n g 18 m [59 feet] long!" T h e authors refer to this creature as
"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
1991). Tarlo (i960) described this arrangement as follows: " T h e symphysis is
extremely short and so expanded that its 5 pairs of large caniniform teeth are
set almost in a circle. I find it difficult to offer any functional explanation of
this remarkable dentition." T h e r e is an explanation, but it is not a functional
one. In his 2001 Ph.D. dissertation, N o e wrote:
W h e n the teeth of Simolestes were believed to have been splayed out, it was
assumed that the animal fed on large vertebrate prey, but with Noe's revision,
"Simolestes is reinterpreted as primarily teuthophagous [squid e a t i n g ] , consum-
ing belemnitcs, soft teuthids or ammonites."
From the H a r r a r region of Ethiopia, von H u e n e (1938) described a frag-
ment of the snout of a pliosaur that he identified as Simolestes nowackianus, but
in 1996, Nathalie Bardet and Stephane H u a wrote that the supposed African
species Simolestes nowackianus is based on part of a jaw belonging to the tele-
osaurid crocodile Machimosaurus. T h e newly described Maresaurus coccai from
Argentina (Gasparini 1997) is also a simolestine pliosaur, with a spatulate,
tooth-studded rosette on the end of its jaws. (It now appears that the spatu-
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 syn-
onymous, 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 cur-
rently 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 reconstruc-
tion 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 frag-
ments of a pectoral girdle —material that many researchers (but not a l l )
consider inadequate to diagnose a genus and species. "A full-grown Megalneu-
saurus m a y have been in the 35—40-foot range or larger," wrote Creisler, "but
such estimates should be greeted with caution until more hard facts are
known."
0
For this paper, Adams also d a w a reconstruction of Trinacromerum bonneri that might have
What does
Trinacromerum
(1993b) wrote that this type of long-snouted pliosaur (called dolichorhyn-
remind you of? A
chopine because of Dolicborbyncbops) became the most common short-necked
10-foot-long, four-
plesiosaurs in the Western Interior Seaway after the extinction of the ich-
flippered penguin? It
was actually one of thyosaurs at the end of the Jurassic period. Indeed, they began to resemble
the last of the 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 charac-
teristic 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 struc-
ture 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 Pachycosta-
saurus 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."
with a short neck and a large head. T h e real evolutionary story may be more to the bottom and fed
complex, and some researchers now think that the pliosaur design may have on mud-dwelling,
developed more than once. Each of the fearsome short-necked forms — soft-bodied prey such
shrimps.
skull with the biting power of killer whales and crocodiles. Exactly how large
Pachycosta-
these great predatory sea dragons grew remains an intriguing question. T h e r e
saurus was about
is no question, however, that these prehistoric reptiles are the quintessential
to feet long.
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.
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 con-
taining 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 myste-
riously 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
everfound—and the
largest, reaching 58
feet. The enormous
size of this animal,
believed it was a toothed whale; French scholar Barthclmy Faujus dc Saint-
along with its
Fond ( k n o w n as Faujus) published an elaborate description in which he
powerful jaws and
compared it with a crocodile. Unimpressed with its size, Pieter Camper's son
teeth, meant that
Adriaan Gillcs C a m p e r correctly compared it with a varanid lizard, and based
almost any creature
was potential prey, on his correspondence with Adriaan Camper, Baron Cuvier opined that le
including hard- grand animal fossile de Maastricht should be placed somewhere between iguanas
shelled turtles and and varanid lizards. " T h e lizard status of mosasaurus," wrote L i n g h a m - S o l i a r
probably even other in 1995, "was first c o m m u n i c a t e d by A. G. C a m p e r in letters to Cuvier in 1790
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
a crocodilian that came from an iron mine in Bavaria, and he eventually
changed the name to Geosaurus ( C a m p 1942). It was not until 1822 that the
animal was awarded a Linnean binomial, when the Reverend Conybeare called
it Mosasaurus — from Mosa, the Latin name of the M a a s ( M e u s e ) River near
Maastricht, and saurus, for "lizard." Later, Dr. Hoffmanns name became a
permanent part of the binomial.*
Theagarten Lingham-Soliar, now of the University of Durban-Westville
in Kwazulu-Natal, South Africa, published a detailed description of Mosa-
saurus hoffmanni in 1995, when he was a Royal Society of L o n d o n research
fellow at the Geological Institute of the University of Tubingen in Germany.
He wrote, "Although the first specimen was described over 200 years ago, it is
here fully described for the first time to provide detailed insights into its
anatomy, functional m o r p h o l o g y and evolution." In accordance with C o p e s
rule (that, over time, there is a general trend toward greater s i z e ) , M. hoffmanni
was one of the latest and largest of the mosasaurs, reaching a length of 58
feet.f Its skull was telescoped, with the nostrils moved back from the end of
the snout, but less so than in the m o d e r n cetaceans. T h e teeth were more
complex than those of any earlier mosasaur species, with multifaceted edges
that made for more effective cutting or breaking of prey items. Its eyes were
large, but its binocular vision was limited, as it was in most mosasaurs except
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. Ac-
cording to L i n g h a m - S o l i a r and N o l f ' s 1989 description of Prognathodon (orig-
inally 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.
("projecting jaw teeth"). T h e rest of its dental a r m a m e n t consisted of large,
ridged jaw teeth and a particularly nasty set of palatal teeth.
T h r o u g h o u t the long history of vertebrate evolution, several unrelated
groups have abandoned the land for an aquatic ( o r s e m i a q u a t i c ) lifestyle.
Cetaceans (whales and d o l p h i n s ) , which are descended from terrestrial q u a d -
rupeds, began their return to the sea about 45 million years ago. W i t h the loss
of their hind legs and the transformation of forelegs into flippers and tail into
flukes, they have achieved a totally aquatic existence. Sirenians (manatees and
d u g o n g s ) accomplished the same thing, and they too lost their hind legs and
acquired flukes (the dugong has a forked tail, and the manatee has a rounded
o n e ) . Pinnipeds (seals, sea lions, w a l r u s e s ) appear to be in a transitional
mode, not yet fully aquatic and spending a large portion of their time on land.
T h e penguins, descended from flying birds, have traded their wings for
flippers and now do all their "flying" underwater. T h e ichthyosaurs are de-
scended from land reptiles of some sort, but we have no idea what the latter
might have looked like —the earliest ichthyosaurs looked much like the later
ichthyosaurs, only less streamlined —but in order to take the plunge, the
plesiosaurs developed such adaptations as pachyostotic gastralia, vcntrallv
located limb girdles, hydrofoil limbs, and specialized ears, eyes, and palates.
Perhaps connected with the demise of the ichthyosaurs, the mosasaurs a p -
peared rather suddenly in the fossil record in about the m i d d l e of the Creta-
ceous, some 90 million years ago in what is now the chalk hills of Kansas, as
well as in Alabama, central Africa, and northern Europe. ( T h e inundation of
their habitat no doubt accelerated the evolution of aquatic lizards, but it
provides no explanation for the disappearance of the ichthyosaurs, their
predecessors as the seaway's dominant predators.)
smallish terrestrial
unlike today's
they had arisen earlier." As Carroll and DeBraga (1992) wrote, " N o significant ancestors of
characters of the skull are known that distinguish aigialosaurs from primitive mosasaurs (and
mosasaurs. In contrast, the trunk vertebrae and limbs arc indistinguishable varanid lizards)
from those of terrestrial varanoids." T h e y are considered a "sister-group" of might have looked
the mosasaurs, which "suggests an earlier Cretaceous dichotomy separating something like this.
advanced varanids and aigialosaurs from the more primitive genera now
included in the Varanidae."
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 an-
cestral mosasauroids, then something very similar began the mosasaur line,
which Gordcn Bell (1997a) called a "27 million—year 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 ani-
mals. 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.)
inhabitant,
* In general usage, osteoporosis is a disease in which bones become increasingly porous, brittle,
and subject to fracture. I'aehyoslosis, which means "thickening of the bone," is the natural
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 pneumatic ("air-filled," like the bones of most birds), tenuiosis
(from tenuis, meaning "thin"), and elaphrosty (from the Greek eiaphro, meaning "light in
weight"), but there is no generally accepted term.
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 past and present, mosasaurs were air breathers.)
Tylosaurus
" T h e seas that rolled over Kansas in Cretaceous times," wrote Charles
proriger (after
Gilmore in 1921, "contained thousands of these a n i m a l s [mosasaurs] and in
Oshorn t8gg).
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 par-
allel 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 col-
lected 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
Western Interior Seaway covered most of midwestern N o r t h A m e r i c a from
the G u l f of M e x i c o to the Arctic Circle, incorporating all of Saskatchewan,
N o r t h and South Dakota, Kansas, Nebraska, O k l a h o m a , and most of Texas.
As of Russell's 1993 summary, the N i o b r a r a C h a l k formations of Kansas had
yielded no fewer than 1,823 mosasaur specimens, the great majority of which
were collected by the O. C. M a r s h and E. D. C o p e expeditions of the late
nineteenth century. Since that time, more have been found in Kansas and
elsewhere, but no other location on Earth has provided so much material for
the study of these marine lizards. Here is Cope's immodest description of one
of his finds, as it appeared in a report prepared for Frederick Hayden's 1871
U.S. Geological Survey of I be Territories:
" Liodon dyspelor 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 Tylosaurtis dyspelor in his list of "Mosasaurs of Uncertain
Taxonomic Position" and wrote, "Tylosaurus dyspelor cannot be referred to cither of the two
species of Niobrara Tylosaurus, and must be regarded as a nomen vanum." He seemed to think
that the T. dyspelor material might belong to Tylosaurus proriger because of its larger size, but he
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 (doubtful name), usually indicating a
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.
* 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 1869—1870
the mosasaurs,
Tylosaurus
proriger lived in
Cretaceous, some 65
preyed on fish,
M i k e Everhart (2002a) reexamined the tylosaur material in the collections shellfish, and
of the Sternberg and other museums, particularly the fossils of Tylosaurus probably smaller
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, and
plcsiosaurs —
T h e first specimens, probably collected by George Sternberg (the older
anything and
brother of Charles), were described by C o p e (1874) and named Liodon nepaeo-
everything that
licus. Cope concluded that T. nepaeolicus was about a third smaller than the more
swam.
common T. proriger but was a separate species and not a juvenile. Everhart
concluded:
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 terres-
trial 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 .
M a r s h described many mosasaurs, including one that lie named —in an
uncharacteristic burst of generosity — Mosasaurus copeanus ( o r perhaps Marsh's
combination of the words Cope and anus was not such a generous act after a l l ) .
Louis Dollo, a Belgian paleontologist, named Plioplatecarpus marshi after M a r s h ,
who had drawn attention to the characters that distinguished it from Mosa-
saurus hofjmanni. T h e n Joseph Lcidy (like Cope, from P h i l a d e l p h i a ) reported
fragmentary mosasaur fossils from N e w Jersey, Alabama, and M i s s i s s i p p i ,
and shortly thereafter, reports began coming in from N e w Zealand, Belgium,
France, and Russia. Around the turn of the century, John C. M e r r i a m de-
scribed many new species from the N i o b r a r a C h a l k of Kansas, including a
second species of Halisaurus, a new species of Platecarpus, and another Ciidastes.
In 1898, Samuel W i l l i s t o n reviewed the known mosasaurs to date and p u b -
lished the results, complete with detailed anatomical drawings, in the Univer-
sity Geological Survey of Kansas. In his 1914 book Water Reptiles of the Past and Present,
W i l l i s t o n wrote:
Although the opening salvos of their lifelong battle were fired over the
misplaced head of the plesiosaur Elasmosaurus in 1869, C o p e and M a r s h re-
mained bitter enemies until death brought an end to their hostilities. ( C o p e
died in 1897; M a r s h two years later.) T h e i r feud escalated in 1871, when C o p e
made an expedition to Kansas, covering ground that M a r s h considered his
exclusive province. T h e n C o p e invaded W y o m i n g , another of Marsh's favorite
hunting grounds, where he found a hitherto unknown trove of Eocene m a m -
mal fossils. It was around this t i m e that M a r s h utilized the newly raised
telegraph lines to relay his discoveries to N e w Haven, and the battle was
Probably an
inhabitant oj
shallow inshore
waters,
I 'lioplatccarpus
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, destroy-
ing 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
nephew), M a r s h could better afford this pitched paleontological battle; C o p e
drove himself to the brink of b a n k r u p t c y trying to compete.
Along with actually finding the fossils (or having them found by someone
in your e m p l o y ) , the question of first publication was the primary battlefield.
N a m i n g an animal (fossil or o t h e r w i s e ) gets your name permanently attached
to it, and Cope and M a r s h tried to bestow their names on as many new
species as possible. T h i s competition often resulted in too-hasty identifica-
tions (or simultaneous identifications of the same fossil, each of them giving
it a different n a m e ) , and as a result, an unrealistically large number of fossil
animals was identified, particularly mosasaurs. In Samuel Williston's 1898
revision of the mosasaurs, he wrote that "four fifths of all the described
species must be abandoned," largely because the differences identified by
Cope and M a r s h in their haste were artifacts of preservation and did not
warrant the erection of a new species. In his extensive 1967 review of American
mosasaurs, Dale Russell wrote, " M a r s h and Cope led field parties into the
Niobrara C h a l k of western Kansas in 1870—71 and soon began describing new
mosasaur material. Far too many species were named; all were inadequately
diagnosed; and the resulting confusion has been a serious handicap to subse-
quent workers."
* 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 cone-
shaped 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 straight-
ened 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.
Everything we know about mosasaurs comes from the analysis of fossils,
but from this material, we can tell quite a lot about the lifestyle of these large
seagoing lizards. T h e size of Tylosaurus, for example, is easy enough to deter-
mine; it ranged in size from 20 to 50 feet, about half of which was tail. H o w
much d i d it weigh? A 17-foot-long great white shark weighs about 3,000
pounds, but it doesn't have a particularly long tail, so a better comparison
m i g h t be the saltwater crocodile (Crocodylus porosus), which has a pretty long
tail and, at a length of 20 feet, weighs about 2,500 pounds. At 50 feet, the
"Bunker" Tylosaurus, found in western Kansas around 1910, was one of the
largest mosasaurs ever found in the U n i t e d States. Its skull was 6 feet long,
and it had 72 sharp, backward-curving teeth in its jaws. It might have weighed
as much as 8 tons.
T h e genus Mosasaurus includes the Maastricht species and probably the first
mosasaur ever found in America. In 1804, Lewis and C l a r k found the remains
of a huge "snake" on an island in the Missouri River in what is now South
Dakota, but the fossil was sent back to Washington and lost. (It was probably
M. missouriensis, which W i l l i s t o n originally named M. borridus.) T h e r e was great
variety among the mosasaurs; some were the size of dolphins, and others were
as big as small whales. T h e round-toothed Clobidens is unlike any other
mosasaur, and Plioplatecarpus marshi may have flown through the water like a
gigantic penguin.
But where? H o l m e s , Caldwell, and C u m b a a (1999) examined a specimen of
Plioplatecarpus found in the Scabby Butte Formation of Alberta and wrote that
"the m o r p h o l o g y of the well-preserved forelimb indicates that the animal
cannot be reconstructed as a subaqueous flyer, but probably used its forelimbs
for paddling. T h e associated matrix . . . suggests that this mosasaur was able
to exploit estuarine and freshwater environments." M o s t mosasaurs were
marine, but this specimen was found in sediments that strongly suggest "an
overbank deposit representing flooding of a coal s w a m p occurring as a lateral
equivalent to the deltaic channel system." T h e geological evidence ("the only
robust diagnostic clues for hypothesizing paleocnvironments") indicates that
this largc-flippered Plioplatecarpus lived inland, perhaps in a freshwater estuary.
W h e n Kenneth W r i g h t and S a m u e l S h a n n o n (1988) found "an unusual
mosasaur" in the uncataloged collections of the University of Alabama M u -
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 spe-
cific 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 ich-
thyosaurs, 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 mosa-
saurs, 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 ad-
vanced embryos distributed along the posterior two-thirds of the trunk re-
gion. T h i s orientation suggests that they were born tail first (the nostrils
emerging l a s t ) to reduce the possibility of drowning, an adaptation shared
with other highly aquatic amniotcs such as cetaceans, sirenians, and ich-
thyosaurs. . . . Viviparity in early medium-sized amphibious aigialosaurs may
have freed them from the need to return to land to deposit eggs, and permit-
ted the subsequent evolution of gigantic totally marine mosasaurs." M i k e
Everhart (2002b) wrote, "Despite problems related to the preservation of
smaller individuals, a review of more recently collected material shows that
immature mosasaurs (estimated body length = 2 m or less) are well repre-
sented throughout the [ N i o b r a r a ] chalk. T h e recent discovery of fetal mate-
rial associated with a mosasaur from South Dakota, and with a mosasaurid
from Slovenia, provide compelling evidence that these marine reptiles bore
live young."
A d d i t i o n a l evidence of sharks scavenging on mosasaur skeletons was un-
earthed near Liege, Belgium, by Bardet et al. (1998). T h e y found caudal
vertebrae identified as belonging to Plioplatecarpus marshi, a mosasaur found in
the Netherlands and Belgium, but very little cranial material, because this
species had a particularly fragile skull. T h e vertebrae had "elongate and
slender grooves on the neural spine, which arc here interpreted to be the result
of scavenging by the c o m m o n dogfish ( s q u a l i d ) shark, Ccntropborodes appen-
diculatus." ( T h i s shark, whose teeth resembled those of the living dogfish
[Scjiialus], lived during the Maastrichtian stage, as did Plioplatecarpus.) T i m
Tokaryk (1993) of the Saskatchewan M u s e u m of N a t u r a l H i s t o r y wrote that
"interest is resurfacing in N o r t h American mosasaurs, as illustrated by the
work performed by Rothschild and M a r t i n (1987) on avascular necrosis,
discoveries of sub-adults by Bell and Sheldon (1986) and discussion on tooth
m o r p h o l o g y and its relevance to preferred prey by Massare (1987)." T h e fossil
Bell found was a small mosasaur, Plioplatecarpus primaevus, but it was the most
complete specimen ever found in N o r t h America.
In September 1996, M i k e and Pam Everhart removed the skull and seven
cervical vertebrae of a very large mosasaur from an exposure of the S m o k y
H i l l C h a l k called H o r s e t h i e f Canyon in Gove County, Kansas:
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).
In 2002, C a i t l i n Kiernan published a study of "more than 600 mosasaur
specimens from the Tombigbce Sand M e m b e r (Eutaw F o r m a t i o n ) and Selma
g r o u p of west and central Alabama." After examining most of this material,
she recognized "significant stratigraphic segregation among taxa." Because the
g e o l o g y of various areas differed, suggesting that different conditions pre-
vailed during the time of the mosasaurs, certain mosasaur fossils were found
in some areas and not others. Kiernan identified three biostratigraphic zones:
the Tylosaurus acme zone (where Tylosaurus fossils p r e d o m i n a t e d ) , the Clidastes
acme zone, and the Mosasaurus acme zone. Tylosaurus evidently lived in shal-
lower, nearshore waters, while Clidastes lived in deeper water, farther offshore.
Kiernan mentioned a "mostly unprepared" skeleton of 55-foot Mosasaurus —
probably hoffmanni—"hy far the most complete mosasaur (or any other fossil
vertebrate) to come from the Prairie Bluff Chalk." L i n g h a m - S o l i a r (1995),
describing Af. hoffmanni from the Netherlands, wrote that it "lived in fairly
deep nearshore waters of 40—50 m (131—164 feet) depth, with changing tem-
peratures and rich vertebrate and invertebrate life." He also called it "the
largest marine reptile ever known,"* and it is interesting to realize that these
gigantic predators plied the Cretaceous seas of Europe, N o r t h America, and,
now, western Asia. In 2002, Bardet and Tonoglu published a description
of m a x i l l a r y fragments ( w i t h teeth) of a mosasaur that they identified as
M. hoffmanni found in the late Maastrichtian deposits at Kastamonu, northern
Turkey.
* It may not hold this title for long. As discussed on pp. 8 9 — 9 0 , Betsy Nicholls of the Royal
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 duropha-
gous 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.
feet, it was also one 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.
S w i n t o n (1930b) of the British M u s e u m found the first mosasaur in Nigeria,
he named it Mosasaurus nigeriensis (according to L i n g h a m - S o l i a r [1991b], S w i n -
tons material actually consists of a number of mosasaur taxa lumped together
as Mosasaurus nigeriensis). O t t o Z d a n s k y of the Egyptian University published a
paper in 1935 on the mosasaurs of Egypt and other parts of Africa, identifying
ten different species, eight of which were found in Egypt. In 1969—1970,
an expedition from the University of Florence headed for Nigeria in search
of fossils, and in the Goronyo district of S o k o t o State, they hit pay dirt
( A z z a r o l i et al. 1975). T h e Dukamaje Formation was so rich in vertebrate
fossils that they called it the " M o s a s a u r Shales," and they found, described,
* Evidently, Azzaroli et al. erected the genus Goronyosaurus on the basis of "some highly
aberrant features" of the skull, but when Soliar ( 1 9 8 8 ) examined the material, he found their
description incorrect in some respects, "but other characters described by them, plus the
and named the 25-foot-long Goronyosaurus nigeriensis* a new species that resem-
bled 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:
new information added here, vindicates the erection of a new genus, which can be tenta-
tively assigned to the Tylosaurinac." A new subfamily might be warranted here, but so far,
only the single species has been found.
Found in the Upper (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,
later paper (2002b) only to the caniniform teeth and how Goronyosaurus might
Nigeria, the 21-
have used them: "Taken together with skull strengthening features discussed
foot-long
in a functional study of the skull of Goronyosaurus, the enlarged teeth . . . seem
Goronyosaurus
to be a novel mosasaurian development of the dentition and a food trap. Such
was one of the larger
mosasaurs. With features arc more characteristic of some crocodiles and much earlier terrestrial
teeth that fit into carnivores. . .. Goronyosaurus represents an exemplar of a giant, marine reptilian
opposing sockets like predator at the end of the Late Cretaceous."
those of a crocodile, " T h e Iullcmmeden Basin [southwest N i g e r ] provided one of the richest
this reptile was a environments in the world for mosasaur evolution and diversification," wrote
formidable predator.
L i n g h a m - S o l i a r (1998a).* Here he discovered Pluridens walkeri, which had twice
as many teeth as any other mosasaur (Pluridens means "many teeth"), and their
structure suggested that it fed on small fishes and thin-shelled invertebrates,
rather like some of the ichthyosaurs. Somewhere around 65 million years ago,
in the period we call the U p p e r Cretaceous, this many-toothed mosasaur
* Niger and Nigeria are two different countries, although the great Niger River runs
through both. Niger (pronounced Ni-zhcr), directly north of Nigeria, is largely composed
of semiarid lands or Saharan desert. Nigeria has a long coastline on the Gulf of Guinea: it
consists of mangrove swamps near the coast and, farther inland, savannas, rain forests, and
high plains.
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 ex-
amined 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 underwa-
ter, 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-
nathodon currii (after Phil Curry, a famed C a n a d i a n dinosaur paleontologist)
and wrote that it "represents one of the largest mosasaurid skulls ever dis-
covered, rivaled only by the giant tylosaurinc Hainosaurus bemardi, and the
plotosaurine Mosasaurus hoffmanni, which appears to have been subequal in size
to P. currii." T h e y estimated its total length at about 36 feet and wrote that its
powerful skull and teeth suggest that it was "a top predator, which had
become adapted for predominantly hunting large, vertebrate prey, a sugges-
tion further corroborated by the unusually large size of the palatal teeth.
* The rules of scientific nomenclature arc varied and complex. W h e n Bonde and Chris-
tiansen discussed Oronosaurus at a symposium in Copenhagen in 1 9 9 9 , they were fairly
certain that this gigantic mosasaur specimen represented a new genus and felt comfortable
giving it a new name. For the most part, however, a scientific name is officially accepted only
when it is published in a journal, and Oronosaurus did not qualify. In fact, it now turns out
that it isn't a new genus after all. In a letter to me in December 2 0 0 1 , Christiansen wrote:
"The giant is no longer called Oronosaurus. When I did phylogenetic analyses using only the
American prognathodons (and, of course a larger number of other taxa) the specimen
consistently (under various optimizations) came out as the sister taxon to all of the other
mosasaurines. As the sister taxon to all other mosasaurines it had 10 be a new genus.
However, the American prognathodons are different from the Furopean type species /'
solvayi, and when this animal was included in the analysis, results changed. Now our monster
consistently emerged as the sister taxon to P. solvayi, with the American prognathodons
forming successive out-groups to the pair. Thus, our animal is a Prognathodon, albeit an
enormous, and highly derived species —and it very evidently is a new species."
Evidently, P. currii is a superior candidate to the title of marine tyrannosaur
than any of the other large mosasaurids."
Although the first N e w Zealand plcsiosaur fossils were uncovered in 1859,
recent discoveries have put this isolated island group right in the m i d d l e of
marine reptile research. Ichthyosaur fossils were found in 1954, at M a n -
gahouanga on N o r t h Island. A m a t e u r collector Joan Wiffen (affectionately
known as the "Dragon L a d y " ) has found the remains of plesiosaurs, sharks,
and turtles and several large fragments and skulls of various mosasaurs,
including Tylosaurus baumuriensis, Mosasaurus mokoroa, and a new species she
discovered and named — Moanasaurus mangabouangae. M u c h of this material
is now being studied; the most recent description (Bell et al. 1999) can be
found in the abstracts from the S o c i e t y of Vertebrate Paleontology's annual
meeting:
have been jound in 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 occa-
sionally 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 ich-
thyosaur 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 Glo-
bidens 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 swal-
lowed 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 Pem-
bina 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 mecha-
nism, 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 per-
petrator was Hainosaurus bemardi. Examining a cast of the braincase of M. hoff-
manni 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-
T h e initial attack was directed at the upper side of the conch, providing
the ammonite was s w i m m i n g in the n o r m a l position used by its living
relative, Nautilus. T h e mosasaur must have dived at it from above to seize it.
T h e attack was pertinaceous, resulting in at least sixteen bites. As shown by
the marks of the pterygoid teeth in the sequence of bites, the mosasaur
evidently tried to swallow the entire ammonite, pulling it as far back into
the throat as possible.
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:
* 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 even-
tually 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:
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, "be-
cause 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
similar in shape ( a n d probably in habits) to today's great white and mako
sharks.
Everhart (1999) identified the shark-bitten mosasaurs of the seaway as
Tylosaurus, Platecarpus, and Ciidastes. A large Plioplatecarpus mosasaur from the
Cretaceous Pierre Shale in Griggs County, N o r t h Dakota, was found in 1995
by two local fossd collectors, M i k e Hanson and Dennis Halvorson. T h e y
contacted John Hoganson, the paleontologist for the N o r t h Dakota Geolog-
ical Survey, and spent almost two years excavating the fossil. T h e specimen is
around 70 percent complete, missing only the flippers, pelvis, a few ribs, and
parts of the tail. It is the largest Plioplatecarpus ever found, 25 percent larger than
the next largest, and a new species. T h e skeleton is 23 feet long; the near-
complete skull alone is 3 feet long. Like the Kansas mosasaurs, this fossil
showed numerous shark teeth embedded in the bone, leading to the specula-
tion that this mosasaur had either been killed by sharks or scavenged as it
floated.
I he relationship of mosasaurs to other reptiles was long a subject of
controversy, but Dale Russell's 1967 study, "Systematica and M o r p h o l o g y of
the American Mosasaurs," alleviated much of the confusion. Still, according
to Gordcn Bell (whose 1997 s u m m a r y I have abridged here), "although many
authors have favored varanid-mosasaurid relationships, such a hypothesis has
not been rigorously tested using modern phylogcnetic methods." But, writes
Bell, "as for the relationships among mosasaurids, the picture is much clearer.
M a n y of the relationships proposed by Russell (1967) have been supported by
two fairly vigorous phylogcnetic analyses using many characters." Carroll
(1988) included aigialosaurs and mosasaurs in his discussion of "aquatic
varanoid lizards" and said that Opetiosaurus (an a i g i a l o s a u r ) was about 3 A feet
l
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.
Tylosaurus
attacking a smaller
mosasaur. Ranging
in size from 10 to
predators of the
the Cretaceous
period.
slicers, snappers, piercers, graspers, rammers and crunchers — was unequaled
by other marine reptiles. T h e y were among the most adaptable of animals,
and clearly flourished during this period of changing temperatures and reced-
ing sea-levels. So what went wrong?" According to Dale Russell (personal
c o m m u n i c a t i o n 1999):
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.
tod,n.
T h e n Lee, Caldwell, and Scanlon (1999) reexamined Pachyophis woodwardi,
which had originally been described as a snake by Nopsca in 1923, but "the
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 paleoenviron-
ment and paleoecology of the marine snakes Pacbyopbis and Pacbyrbachis and the
terrestrial snake Dinilysia. As mentioned earlier (Scanlon et al. 1999), Pacby-
rbachis 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 proba-
bly 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 conclu-
sions 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
snakes and concluded that all snakes were unrelated to mosasaurs. T h i s is a
spurious claim as no other lizards of any k i n d were included in their anal-
ysis. . . . Tchcrnov and his collaborators have no replacement hypothesis for
the sister g r o u p relationship of snakes —only that Caldwell and Lee are
wrong. T h a t sort of statement is not science."
In the essay that introduced the Tchcrnov paper in Science, Greene and
C u n d a l l criticized the approach ( a n d conclusions) of Caldwell and Lee
(1997), who "showed their drawings and reconstructions of Pachyrhachis to a
number of nonscicntists who use 'snake' in the vernacular sense, and all
identified Pachyrhachis as a snake rather than a lizard." Even though it had legs,
the dense, heavy bones of Pachyrhachis suggest that it was a water dweller, and
the location of the fossil in marine deposits seems to confirm this suggestion.
Because of similarities in the jaw structure, Lee et al. (1999) concluded that it
( a n d all other snakes) had evolved from mosasaurs. In remarks published in
New Scientist ( H e c h t 2 0 0 0 ) , M i k e Caldwell said, "Rieppcl's analysis fails to
compare the legged snakes with mosasaurs, their closest relatives." In 1998,
H u s s a m Zahcr published a revision of the phylogenetic position of Pachy-
rhachis, and two years later, Caldwell ( 2 0 0 2 b ) responded by writing:
M i c h a e l Lcc believes that the mosasaurs are the nearest relatives of snakes.
Like snakes, mosasaurs had pterygoid palatal teeth and hinged lower jaws.
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 typi-
cal 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 evolu-
tionary origins of snakes. T h e y wrote:
T h e y said that according to Caldwell and Lee, Pachyrhachis is the most primi-
tive 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 phy-
logcny 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-bisection-
reconnection 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:
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 end-
Cretaceous 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.
The "Reason" for Extinction
At one time, the reason for the extinction of the ( n o n a v i a n ) dinosaurs was
regarded as unknown and probably unknowable. Dinosaurs roamed the earth,
and then they didn't. George Gaylord Simpson, writing in 1953, said, " N o one
knows exactly why the dinosaurs and a host of other ancient forms became
extinct. T h i s is not because there is anything mysterious or metaphysical
about extinction or because the possible causes are unknown. It is just because
there are many reasonable possibilities, and the record does not enable us to
say in the particular case which of them were actually involved. All we can say
is that something changed and the dinosaurs did not." In 1968, in The Evidence of
Evolution, Smithsonian Institution paleontologist Nicholas H o t t o n wrote, "It
is sometimes assumed that a sudden major change in climate at the end of the
Mesozoic era was the chief cause of the dinosaurs' demise. However, other
episodes of climactic change during the M e s o z o i c do not appear to have
bothered the dinosaurs excessively, and the suddenness of dinosaur extinction
may be more apparent than real."
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 erup-
tion 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 perma-
nent 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
shocked q u a r t z ( a n indicator of high-temperature i m p a c t ) a n d iridium-laden
layers have now been found around the world at the corresponding strati-
graphic levels. T h e r e was certainly an impact at Chicxulub, and because it
coincided with the mass extinction of the dinosaurs and m a n y other k i n d s of
animals, it is difficult to deny a relationship between the two events, but the
nature of that connection remains unresolved.
W h e t h e r something killed them or they died of unknown causes, the great
terrestrial dinosaurs were all gone after the dust cleared from whatever it was
that happened 65 million years ago. C o n t r a r y to popular conceptions, all the
dinosaurs were not marching around as the Cretaceous came to its abrupt
halt. By that time, most of them were extinct already, and many had been
extinct for tens or even hundreds of m i l l i o n s of years. For example, the
carnivore Allosaurus was gone 135 m i l l i o n years ago, as was Apatosaurus, the great
sauropod that used to be k n o w n as Brontosaurus. T h e feathered reptile Archaecp-
teryx, originally found in shales from 150 million years ago, was gone 30 million
years later, and Iguanodon, the first known dinosaur, vanished no m i l l i o n years
ago. Deinonychus, the dinosaur described as w a r m - b l o o d e d by John O s t r o m in
1969, disappeared from the fossil record 100 m i l l i o n years ago, and Protoceratops,
the predominant dinosaur of the Gobi Desert, was last recorded from strata
dated at 75 million years old. In fact, as far as we can tell from the scanty fossil
record, almost all the dinosaurs had died out except for those in western
N o r t h America. Saltosaurus, Ankylosaurus, Pachycephalosaurus, GaUomimus, 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.
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, sala-
manders, 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.
Still, it is hard to imagine why the ruling reptiles of the sea did not endure.
From the fossil evidence, we can sec that they were numerous, efficient, and
highly diversified. Cetaceans had not yet arrived on the scene, so there was no
competition from them, and the only other large predators were sharks.
Animals can become extinct w i t h o u t a catastrophe, and the vast majority of
the millions of extinctions that have occurred in the geological past have been
caused by a combination of factors whose synergistic influences we simply do
not understand. Groups of animals rise and fall in cycles that arc largely
unknown and unpredictable, but m a n y extinctions were probably related to
global change as the continents slid around, m o u n t a i n s were pushed up, and
oceans cooled, warmed, dried up, or were formed anew. Tectonic changes are
certainly responsible for massive climatic modifications, but we can only
wonder what, if any, effect these earth movements may have had on the lives
of animals.
Adams, D. A. 1 9 9 7 . Trinacromerum bonneri, new species, (Rbomaleosaurus thomtoni, sp. n.) from the Upper
last and fastest pliosaur of the Western Interior Lias of Northamptonshire. Ann. Mag. Nat. Hist.
Seaway. Texas Jour. Sci. 4 9 ( 3 ) 1 1 7 9 — 9 8 . 9('°) 4°7-'5-
:
from the Oxford Clay of Peterborough. Ann. Mag. Aronson, R. 1 9 9 0 . Rise and fall of life at sea. New
Nat. Hist. 4 : 4 1 8 — 2 9 . Scientist 1 2 8 ( 1 7 4 1 ) : 3 4 — 3 7 .
. 1910. A Descriptive Catalogue of the Marine Reptiles of Augusta, ] . , and Z. Burian. 1 9 6 4 . Saurier der Urmeere.
the Oxford Clay, Based on the Leeds Collection in the Artia.
British Museum (Natural History), London, Part I. Azzaroli, A., C. dc Giuli, G. Ficcarelli, and D. Torre.
British Museum. 1 9 7 2 . An aberrant mosasaur from the Upper
. 1 9 1 2 . Description of a new plesiosaur Cretaceous of north-western Nigeria. Mmiorif
(Plesiosaurus capensis, sp. nov.) from the Uitenhage Accademia nazionale dei Lined. Classe di scienze fsche
Beds of Cape Colony. Ann. S. Afr. Mus. 7 : 3 0 9 — 2 2 . malematiche e naturali 5zf j):5j—56.
. 1 9 2 2 a . Description of a new plesiosaur from the . 1 9 7 5 . Late Cretaceous mosasaurs from the
Weald Clay of Berwick (Sussex). Q Jour. Geol. Soe. Sokoto district, Nigeria. Memoric Accademia nazionale
London 7 8 : 2 8 5 — 9 8 . dei Lincei. Classe di scienze fsche mateniatiche e naturali
ichthyosaur from the Upper Jurassic lithographic . 1 8 9 0 . On the characters and systematic position
limestones of Bavaria.your. Paleo. 7 4 ( 3 ) 1 5 0 3 — 1 1 . of the large sea-lizards, Mosasaundae. Science
Bardet, N., and S. Hua. 1 9 9 6 . Simolestes nowackianus 16(405)262.
Huene, 1 9 3 8 from the Upper |urassic of Ethiopia Beaumont, C, G. M. Quinlan, and G. S. Stockmal.
is a telcosaurid crocodile, not a pliosaur. N. fh. 1 9 9 3 . The evolution of the Western Interior Basin:
Ccol Palaeonl. Mh. 1 9 9 6 ( 2 ) 6 5 — 7 1 . Causes, consequences and unsolved problems. In
Bardet, N„ and X. P. Subcrbiola. 2 0 0 1 . T h e basal W. G. E. Caldwell and E. G. Kaufmann, eds.,
mosasaurid Halisaurus sternhergii from the late Evolution of the Western Interior Basin, pp. 9 7 — 1 1 7 .
Texas. In V L. Santucci and L. McClelland, eds., . 1 9 9 3 . Late Triassic extinctions and the origin of
National Park Service Paleontological Research, vol. 2. the dinosaurs. Science 2 6 0 : 7 6 9 — 7 0 .
and ]. McRec. 1 9 9 9 . Sea monsters of the South Benton, M . J., M . A. Wills, and R. Hitchin. 2 0 0 0 .
Pacific: On the Late Cretaceous mosasaurs of Quality of the fossil record through time. Nature
New Zealand.your. Vert. Paleo. I9(suppl. to 3 ) : 3 2 A . 4°3 534-3 -
: 6
Buel, J. W. 1 8 8 7 . Sea and Ijind: An Illustrated History of the . 1 9 9 9 a . Description and phylogenetic
Wonderful and Curious Things of Nature Existing before relationships of a new species of Coniasaurus
and since the Deluge. A Natural History of the Sea, Land Owen, 1 8 5 0 (Squamata).your: Vert. Paleo. 1 9 : 4 3 8 — 5 5 .
Creatures, the Cannibals and Wild Races of the World. . 1 9 9 9 b . Squamate phylogeny and the
Crocodiles and Alligators, pp. 1 4 — 2 5 . Facts on File. . 2 0 0 0 a . On the aquatic squamate Dolichosaurus
Cadbury, D. 2 0 0 0 . The Dinosaur Hunters: A True Story of . 2 0 0 2 . From fins to limbs to fins: Limb
Scientific Rivalry and the Discovery of the Prehistoric World. evolution in fossil marine reptiles. Amer. four. Med.
Fourth Estate. Genet. 9 9 9 9 : 1 — 1 5 .
(?Santonian) of east-central Peru, and the Shastasauridae) from the Upper Triassic of the
taxonomic utility of mosasaur cervical vertebrae. El Antimonio district, northwestern Sonora,
four. Vert. Paleo. I 5 ( j ) : 5 $ 2 — 4 4 . Mexico.your: Paleo. 6 3 ( 6 ) : 9 3 0 — 3 9 .
Coniasaurus crassidens Owen 1 8 5 0 ( S q u a m a t a ) from . 1951. Plotosaurus, a new generic name for
the English Chalk (Cretaceous, Cenomanian). Kolposaurus Camp, preoccupied. Jour. Paleo.
. 2 0 0 1 . Live birth in Cactaceous marine lizards ('arev. I*. ( J. 1 9 7 3 . Fish with warm bodies. Scientific
(mosasauroids). Proc. Roy. Soc. London, Biol. Sei. American 228(2): 36—44.
Evolution of the Western Interior Basin. Geological pp. 2 3 2 — 4 3 . Byron Preiss Visual.
Association of Canada. . 1 9 9 6 . A review of short-necked plesiosaurs
Callaway, J. M . 1 9 9 7 a . Ichthyosauria. In J. M . from the Cretaceous of the Western Interior,
Callaway and E. L. Nicholls, eds., Ancient Marine North America. N. Jb. Geol. Palaeont. Abh.
Reptiles, pp. 3—16. Academic Press. 201(2)1259—87.
system: Documenting an evolutionary constraint. Christiansen, P., and N. Bondc. 1 9 9 9 . Anatomy and
American Zoologist 31:644—54. phylogenetic affinities of a gigantic mosasaur from
Carroll, R. L. 1 9 8 1 . Plesiosaur ancestors from the Israel. In E. Hoch and A. K. Brantsen, eds.,
Upper Permian of Madagascar. Phil. Trans. Roy. Soc. Secondary Adaptation to Life in Water, p. 1 1 . University
Trans. Roy. Soc. London, ser. B, 3 0 9 : 3 4 3 — 9 3 . Coates, M. I., and J. A. Clack. 1 9 9 0 . Polydactyly in
Carte, A., and W. H. Baily. 1 8 6 3 . Description of a new the earliest known tetrapod limbs. Nature 3 4 7 : 6 6 —
species of Plesiosaurus, from the Lias, near Whitby, 69.
Yorkshire.your Roy. Dublin Soc. 4 : 1 6 0 — 7 0 . Coates, M . I., and M . Ruta. 2 0 0 0 . Nice snake, shame
Chaloncr, W. G., and A. Hallam, eds. 1 9 8 9 . Evolution about the legs. Trends in Ecology and Evolution
and the last of the dinosaurs. Phil. Trans. Roy. Soc. Life Books.
London B 5 2 5 : ^ 8 7 — 4 0 0 . Colbert, E. H. 1 9 4 9 . A new Cretaceous plesiosaur
. 1 9 9 } . Disaster theories of dinosaur extinction. from Venezuela. Amer. Mus. Novitates 1 4 2 0 : 1 — 2 2 .
names for preoccupied Walkena Chatterjce, 1 8 9 7 . 1963. Dinosaurs: Their Discovery and I heir lie//,/.
from the Upper Cretaceous of Antarctica. Ceol. Colbert, E. H., R. B. Cowles, and C. M. Bogert. 1 9 4 6 .
Soc. London Spec. Pub. 4 7 : 1 9 7 — 2 1 5 . Temperature tolerances in the American alligator
Chatterjee, S., and W. J. Zinsmeister. 1 9 8 2 . Late and their bearing on the habits, evolution, and
extinction of the dinosaurs. Bull. Amer. Mus. Nat. . 1 8 7 0 b . On Elasmosaurus platyurus Cope. Amer. Jour.
Hist. 8 6 : 3 3 1 - 7 3 . Sci. 5 o ( i 4 8 ) : i 4 o — 4 1 .
Coles, H . 185^. On the skin of the Ichthyosaurus. Proc. . 1 8 7 0 c . On some reptilia of the Cretaceous
Ceol. Soc London 9 ( 1 ) 1 7 9 — 8 1 . formation of the United States. Proc. Amer. Phil.
Collin, R., and C. M. Jam's. 1 9 9 7 . Morphological Soc. 11:271—74.
eds., Ancient Marine Reptiles, pp. 4 5 1 — 6 6 . Academic . 1 8 7 1 b . On the geology and paleontology o f the
Cope, E. D. 1 8 6 7 . T h e fossil reptiles of New Jersey. . 1 8 7 2 a . On Kansas vertebrate fossils. Amer. four.
. 1 8 6 8 a . A large new cnaliosaur. Proc. Acad. Nat. Sci. . 1 8 7 2 b . [On the structure of the Pythono-
. 1 8 6 8 b . On a new large cnaliosaur. Amer. Jour. Sci. . 1 8 7 3 . Catalogue of the Pythonomorpha found
4 0
6
57)^63-64. in the Cretaceous strata of Kansas. Proc. Amer. Phil.
. 1 8 6 8 c . On some Cretaceous reptilia. Proc. Acad. Soc. 1 2 : 2 6 4 — 8 7 .
Reptilia) from the Lower Lias (Hettangian: Darwin, C. 1 8 5 9 . On the Origin of Species. John Murray.
Lower Jurassic) of Lyme Regis, England: A Dawson, J . W. 1 9 0 3 . The Story of Earth and Man. Harper
Phosaurid-plesiosaurid intermediate? Zool. jour. and Brothers.
Linn. Soe. 1 1 2 : 1 5 1 — 7 8 . DeBraga, M., and R. L. Carroll. 1 9 9 3 . T h e origin of
. 1 9 9 7 . An early Cretaceous pliosauroid from mosasaurs as a model of macro-evolutionary
South A f r i c a . Ann. S. Afr. Mus. 1 0 5 : 2 0 7 — 2 6 . patterns and processes. £W. Biol. 2 7 : 2 4 5 — 3 2 2 .
Sauropterygia). Ree. W. Aust. Mus. Suppl. 5 7 : 2 0 1 — 5 . Delair, J. B., and W. A. S. Sarjcant. 1 9 7 5 . Joseph
Cruickshank, A. R. I., D. M. M a r t i l l , and L. F. Noe. Pentland — a forgotten pioneer in the osteology of
1 9 9 6 . A pliosaur (Reptilia, Sauropterygia) marine reptiles. Proc. Dorset Nat. Hist. Archaeol. Soc.
exhibiting pachyostosis from the middle Jurassic 1975:12—16.
o f England. Jour. Ceol. Soc. London 1 5 3 : 8 7 3 — 7 9 . Desmond, A. J. 1 9 7 6 . The tlot-Blooded Dinosaurs. Dial
Cruickshank, A. R. I., P. G. Small, and M. A. Taylor. Press.
1 9 9 1 . Dorsal nostrils and hydrodynamically driven . 1982. Archetypes and Ancestors: Palaeontology in
Victorian London 1850—l8jj. University of Chicago Druckenmiller, P. S. 2 0 0 2 . Osteology of a new
Press. plesiosaur from the Lower Cretaceous (Albian)
Dietz, L. E, and W. A. S. Sarjeant. 1 9 9 3 . L. B. Thermopolis Shale of Montana. Jour. Vert. Paleo.
Halstead: A bibliography of his published 22(l):2 - 2.
9 4
Dinosaur Evolution and the Origin of Birds. Freeman. Du Toit, A. I. 1 9 2 7 . A geological comparison of
Dingus, L„ E. S. Gaffncy, M. A. Norell, and S. D. South America with South Africa. Carnegie Inst.
Sampson. 1 9 9 5 . The Halls of Dinosaurs: A Guide to Wash. Pub! 381:1-157.
. 1 8 8 9 . Premiere note sur les mosasauriens de Edinger, T. 1 9 3 5 . Pistosaurus. N. fb. Geo! Palaeont.
Mesvin. Bull. Soc. Belg. Geo! Paleont. Hydro! 3 : 2 7 1 — (74)321-59.
Maastrichtian type area.your. Vert. Paleo. 2i(suppl. Etheridge, R. 1 8 8 8 . On additional evidence of the
to 3):45A-46A. occurrence of Plesiosaurus in the Mesozoic rocks of
Dortangs, R. W, A. S. Schulp, E. W. A. Mulder, Queensland. Proc. Linn. Soc. N.S.W2:410—13.
Formation of western Kansas. Trans. Kansas Acad. . 1 9 9 9 . A new ichthyosaur from the Los Mollcs
Sci. 105(1-2)33-43. Formation (early Bajocian), Neuquen Basin.
. 2002b. Remains of immature mosasaurs Argentina, your Paleo. 7 3 ( 4 ) 6 7 7 — 8 1 .
(Squamata, Mosasauridac) from the Niobrara . 2 0 0 0 . Late Jurassic ichthyosaurs from the
Formation (late Cretaceous) argue against Neuquen Basin, Argentina. Historical Biology 14:133—
4 ): 2A.
5 . 2 0 0 1 . Dorsal or ventral? Homologies of the
. 2002c. W h e r e the elasmosaurs roam. Prehistoric forcfin of Caypullisaurus (Ichthyosauria:
Times 53:24—27. Ophthalmosauria). Jour. Vert. Paleo. 2 1 ( 3 ) 1 5 1 5 — 2 0 .
Everhart, M. J., and J. Bussen. 2001. First report of Fernandez, M . S., and M . Iturraldc-Vinent. 2 0 0 0 . An
the mosasaur, Plioplatecarpus cf. primaevus, from the Oxfordian Ichthyosauria (Reptilia) from Vinalcs,
Pierre Shale ( C a m p a n i a n ) of western Kansas. western Cuba: Paleobiogcographic significance.
Kansas Acad. Sci. Trans. 2o(abstracts):28—29. Jour. Vert. Paleo. 2 o ( i ) : i 9 i — 9 3 .
Everhart, M. J., and P. A. Everhart. 1996. First report Fish, F. E., and J. M. Battle. 1 9 9 5 . Hvdrodynamic
of the shell crushing mosasaur, Globidens sp., from design of the humpback whale flipper.your.
the Sharon Springs Member of the Pierre Shale Morphol. 2 2 5 : 5 1 — 6 0 .
Fleming, C. A., D. R. Gregg, and S. P. Welles. 1 9 7 1 . . 1998. The Broadhill Swordfishery in the North Atlantic.
Anatomy and histology of plcsiosaur hones from . 1 9 2 7 . Note on a second occurrence of the
the late Cretaceous of Seymour Island. Antarctic mosasaunan reptile Globidens. Science 6 6 : 4 5 2 .
Frey, E., and J. Riess. 1 9 8 2 . Considerations Gould, S. G., and N. Eldrcdge. 1 9 7 7 . Punctuated
concerning plcsiosaur locomotion. N. Jb. Geol. equilibria: T h e tempo and mode of evolution
Palaeont. 164:193—94. reconsidered. Paleobiology 3:115—51.
Frey, E., M . - C . Buchy, and W. Stinncsbeck. 2 0 0 1 . T h e Gould, S. J. 1 9 9 0 . Bent out of shape. Natural History
monster of Arambcrn and friends: New finds of 5(90)12-25.
. 1 9 9 7 . A new pliosaur from the Bajocian of the . 1 9 8 6 . End-Cretaceous mass extinction event:
Neuquen Basin, Argentina. Palaeontology 4 0 : 1 3 5 — 3 7 . Argument for terrestrial causation. Science
Gasparini, Z., and L. Spalletti. 1 9 9 3 . First Callovian 238:1237-42.
plesiosaurs from the Neuquen Basin, Argentina. . 1 9 8 9 . T h e case for sea-level change as a
Ameghiniana 3 0 ( 3 ) 2 4 5 — 5 4 . dominant causal factor in mass extinction of
Gasparini, Z., N. Bardct, and M. Iturraldc-Vinent. marine invertebrates. Phil. Trans. Roy. Soc. London
Oxfordian (late Jurassic) of Cuba. Geohios 35:201— Halstead, L. B. 1 9 6 8 . The Pattern of Vertebrate Evolution.
11. Freeman.
Gibson, C. D. 1 9 8 1 . A history of the swordfishery in . 1 9 7 1 . Liopleurodon rossicus (Novozhilov) — a
pliosaur from the Lower Volgian of the Moscow . 1 9 9 3 . Tail-bends of Triassic ichthyosaurs: A re-
Basin. Palaeontology 14:566—70. appraisal. Jour. Vert. Paleo. 1 ^(^):4iA.
. 1 9 8 2 . The Search for the Past. Doublcday. . 1 9 9 4 . Speculations on the role of marine reptile
. 1 9 8 9 . Plesiosaur locomotion. Jour. Ceol. Soc. deadfalls in Mcsozoic deep-sea paleoecology.
London 1 4 6 : 3 7 — 4 0 . Palaios 9 ( 1 ) 4 2 — 4 7 .
. 1 8 3 4 b . Notice of fossil bones found in the Holmes, R., and H.-D. Sues. 2 0 0 0 . A partial skeleton
Tertiary formation of the state of Louisiana. of the basal mosasaur Halisaurus platyspondylus from
Trans. Amer. Phil. Soc. 4 : 3 9 7 — 4 0 3 . the Severn Formation ( U p p e r Cretaceous:
. 1 8 4 2 . On the discovery of the Basilosaurus and Maastrichtian) of M a r y l a n d . Jour. Paleo. 7 4 ( 2 ) ^ 0 9 —
Hawkins, T. 1 8 3 4 . Memoirs of Ichthyosauri and Plesiosauri, Holmes, R., M. W. Caldwell, and S. L. Cumbaa.
Extinct Monsters of the Ancient Earth. Relfc and 1 9 9 9 . A new specimen o f Plioplatecarpus
Fletcher. (Mosasauridac) from the Lower Maastrichtian of
Hcatwolc, H. 1 9 9 9 . Sea Snakes. Krieger. Alberta: Comments on allometry, functional
Hccht, J. 2 0 0 0 . Prehistoric pins. New Scientist 1 6 5 : 1 5 . morphology, and paleoecology. Can. Jour. Earth Set.
Herald. E. S. 1 9 6 1 . Living Fishes of the World. Doublcday. 36:363-69.
Heyning, J. E., and J. G. Mead. 1 9 9 6 . Suction feeding Home, E. 1 8 1 4 . Some account of the fossil remains
in beaked whales: Morphological and more nearly allied to fishes than any other classes
observational evidence. Contrih. Sci. L.A. County of animals. Phil. Trans. Roy. Soc. London 1 0 4 : 5 7 1 — 7 7 .
extinctions? Phil. Trans. Roy. Soc. London B 3 2 5 : 1 3 — 2 1 . . 1 8 1 9 a . An account of the fossil skeleton of the
Hoglcr, J. A. 1 9 9 2 . Taphonomy and paleoecology of Proteosaurus. Phil. Trans. Roy. Soc. London 1 0 9 : 2 0 9 —
Ichthyosaurs: A History of Fossil Sea-Dragons. National Kear, B. P. In press. A juvenile pliosaur from the early
Museum of Wales. Creraceous of South Australia. Lethaia.
Humphries, S., and G. D. Ruxton. 2 0 0 2 . W h y did Kiernan, C. R. 2 0 0 2 . Stratigraphic distribution and
some ichthyosaurs have such large eyes? four. Exp. habitat segregation of mosasaurs in the Upper
Biol. 2 0 5 : 4 3 9 — 4 1 . Cretaceous of western and central Alabama, with
Hungerbiihlcr, A. 1 9 9 4 . Recently identified type an historical review of the Alabama mosasaur
material of the Lower Jurassic ichthyosaur discoveries. Jour. Vert. Paleo. 2 2 ( 1 ) 9 1 — 1 0 3 .
Netherlands, four. Vert. Paleo. icj(suppl. to 3 ) 5 5 . Langton, J. 1 9 9 4 . "Revealed: T h e Loch Ness picture
Kase, T, P. Johnston, A. Seilacher, and J. B. Boyce. hoax. Monster was a toy submarine." Sunday
1 9 9 8 . Alleged mosasaur bite marks on late Telegraph, March 13, pp. 1, 3.
Cretaceous ammonites arc limpet Lee, M. S. Y. 1 9 9 7 . T h e phylogeny of varanoid lizards
(patellogastropod) home scars. Geology and the affinities of snakes. Phil. Trans. Roy. Soc.
26(10)947-50. London B 3 5 2 : 5 6 — 9 1 .
Kass, M., and D. Smith. 2 0 0 1 . Preliminary report on . 1 9 9 8 a . Anatomy and relationships of Pachyrhachis
new material for Prognathodon stadtmani. Jour. Vert. problematicus, a primitive snake with hind limbs.
Paleo. 2i(suppl. to 3 ) 6 7 A . Phil. Trans. Roy. Soc. London 3 5 3 ( 1 5 7 5 ) 1 5 2 1 — 2 2 .
Bioehronology of North America, pp. 2 7 3 — 3 0 6 . Gcol. . 2 0 0 0 . Snake origins and the need for scientific
Soc. Canada Spec. Paper 2 7 . agreement on vernacular names. Pakobiology
Kauffrnan, E. G., and W. G. E. Caldwell. 1 9 9 3 . T h e 27(1)1-6.
Western Interior Basin in space and time. In Lee, M. S. Y, and M. W. Caldwell. 1 9 9 8 . Anatomy
W. G. E. Caldwell and E. G. Kaufmann, eds., and relationships of Pachyrhachis problematicus, a
primitive snake with hmdlimbs. Phil. Trans. Roy. Soc. . 1 9 9 3 . I he mosasaur Lciodon bares i t s teeth.
London B 3 5 3 : 1 5 2 1 — 5 2 . Modern Geology 18:443—58.
1 9 9 9 . A second primitive marine snake: Pachyophis (Reptilia: Mosasauridac) from the Turanian of
woodwardi from the Cretaceous of Bosnia- Angola re-interpreted as the earliest member of
Herzegovina. Jour: Zool. London 2 4 8 : 5 0 9 — 2 0 . the genus Platcearpus. Palaontol. Z. 6 8 ( 1 — 2 ) 2 6 7 — 8 2 .
Li, C , and H.-L. You. 2 0 0 2 . Cymbospondylus from the . 1 9 9 8 b . Iaphonomic evidence for fast tuna-like
Upper Triassic of Guizhou, China. Vertebrata swimming in Jurassic and Cretaceous ichthyosaurs.
PalAsiatiea 4o(i):9—16. N. Jb. Geol. Palaeont. Abh. 2 0 7 ( 2 ) : i 7 i — 8 3 .
Li, J.-L., ]. Liu, and O. Rieppel. 2 0 0 2 . A new species . 1 9 9 8 c . Unusual death of a Cretaceous giant.
of Lariosaurus (Sauropterygia: Nothosauridae) I.ethaia 31:308—10.
flew through the Mesozoic seas — but probably on Queensland pliosaur. Mem. Old. Mus. 1 0 : 1 — 7 .
two wings rather than four. Guardian, November 16. . 1932. Restoration of Kronosaurus tjucenslandicus.
i Lepidosauromorpha, Squamata) from the Upper . 1 9 3 5 . Palacontological notes. Mem. Qld. Mus.
Cretaceous of Belgium and the Netherlands. Paleo. 10:236—39.
Long, ]. A. 1 9 9 5 . The Rise of Pishes: jOO Million Years of . 2 0 0 1 . Observations on Triassic ichthyosaurs.
Evolution. Johns Hopkins University Press. Part VII. New data on the osteology of
. 1 9 9 8 . Dinosaurs oj Australia and New Zealand, and Chaohusaurusgeishanensis Young & Dong, 1972 from
Other Animals of the Mesozoic Era. Harvard University the Lower Triassic of Anhui, China. N. Jb. Geol.
Press. Palaeont. Abh. 2 1 9 ( 3 ) 3 0 5 — 2 7 .
middle Triassic of the Germanic Basin. N. Jb. Geol. . 1 8 8 0 . New characters of mosasauroid reptiles.
Palaeont. Abb. 209:105—34. Amer. Jour. Sci. 3 : 8 3 — 8 7 .
Triassic ichthyosaur Conteetopalatus from Germany. . 1 9 9 5 . An ichthyosaur with well preserved soft
Palaeontology 44(6)1127—56. tissue from the Sinemurian of southern England.
. 2 0 0 1 b . Observations on Triassic ichthyosaurs. Palaeontology 38:897—903.
Part VIII. A redescription of Phalarodon major (von Martill, D. M., and J. D. Hudson, eds. 1 9 9 1 . Fossils of
Huene, 1 9 1 6 ) and the composition and phylogeny the Oxford Clay. Palacontological Association.
of"the Mixosauridae. N. Jb. Geol. Palaeont. Abb. M a r t i l l , D. M., and D. Naish. 2 0 0 0 . Walking with
. 1 8 7 2 c . On the structure of the skull and limbs Martin, L. D, and J. D. Stewart. 1 9 7 7 . The oldest
(Turanian) mosasaurs from Kansas. Jour. Paleo. Ross, cd., Crocodiles and Alligators, pp. 4 2 — 5 7 . Facts
5<5>973-75- on File.
. 1 9 8 2 . An ichthyornithiform bird from the M c C l m t o c k , J. 2 0 0 0 . Romancing the bone. Discover
ichthyosaurs: Evidence and implications. Jour. Vert. quadrtseissus (Reptilia, Ichthyosauridac) Life Sci.
Paleo. 8( ):2iA.
5
Contrib. Roy. Ontario Mus. 93:1—21.
. 1 9 9 0 . The affinities and ecology of Triassic . 1 9 7 3 c . A note on the most recent ichthyosaur
ichthyosaurs. Ceol. Soc Amer. Bull. 1 0 2 : 4 0 9 — 1 6 , known: An isolated coracoid from the Upper
. 1 9 9 4 . Cymbospondylus (Ichthyosauria: Campanian of Saskatchewan (Reptilia.
Shastasuridae) from the Lower Triassic Thaynes Ichthyosauria). Can. Jour. Earth Sci. 1 0 : 1 3 4 6 — 4 9 .
«(3) »7-25-
: . 1 9 7 4 b . A revision of the longipinnatc
Mazin, J.-M., V Suteethorn, E. Buffetaut, J.-J. Jaeger, with descriptions of two new species (Reptilia:
and R. Hclmke-Ingavat. 1 9 9 1 . Preliminary Ichthyosauria). Life Sci. Contrib. Roy. Ontario Mus.
Triassic of T h a i l a n d . Comptes Rendus de I'Academic des relationships of a new ichthyosaur genus from the
Sci. I 5 ( i ) : i 6 g — 7 1 .
. 1 9 9 2 b . Unusual extensions of the neural spines
. 1 9 7 9 a . A revision of the Lower Jurassic in two ichthyosaurs from the Lower Jurassic of
ichthyosaurs from Germany with a description of Holzmaden. Can. Jour. Earth Sci. 2 9 ( 2 ) 1 3 8 0 — 8 3 .
two new species. Paleentographica 1 6 6 : 9 3 — 1 3 5 . . 1 9 9 3 . A new species of large, long-snouted
. 1 9 7 9 b . Selection pressure for high body ichthyosaur from the English Lower Lias. Can.
temperatures: Implications for dinosaurs. four. Earth Sci. 3 0 ( 6 ) 1 1 9 7 — 1 2 0 4 .
Paleobiology 5(3)285—95. . 1 9 9 4 a . Diatoms to Dinosaurs: The Size and Scale of
. 1 9 8 3 . The Successful Dragons. Samuel Stevens. Living Things. Penguin.
. 1 9 8 6 . A putative ancestor for the swordfish-likc . 1 9 9 4 b . A new species of Shastasaurus (Reptilia:
ichthyosaur / urhmosaurus. Nature 1,2.2. 6078 454—56. Ichthyosauria) from the Triassic of British
. 1 9 8 8 . Differential development of the rostrum Columbia: T h e most complete exemplar of the
and mandible of the swordfish (Xiphiasglatlius) genus. Jour. I'erl. Paleo. 1 4 : 1 6 8 — 1 7 9 .
during ontogeny and its possible functional . 1 9 9 4 c . T h e taxonomic status of the Upper
significance. Can. ]our. Zool. 6 6 : 4 9 6 — 5 0 3 . Liassic ichthyosaur Eurhinosaurus longirostris.
. 1 9 8 9 a . Computed tomography reveals further Palaeontology 37(4)747-53.
details ot Exealibosaurus, a putative ancestor tor the . K)94d. Tcmnodontosaurus riser is a juvenile form ot
swordfish ichthyosaur Eurhinosaurus. Jour. Vert. Paleo. T. platydon. Jour. Vert. Paleo. 1 4 : 4 7 2 — 7 9 .
9(3)269-81. . 1 9 9 5 a . A new and typically |urassie ichthyosaur
. 1 9 8 9 b . T h e lehthvosaunan tailbend: A trom the Upper Triassic of British Columbia.
verification problem facilitated by computer Cm. Jour. Earth Sci. 3 3 : 2 4 — 3 2 .
tomography. Paleobiology 15(4)1429—36. . 1 9 9 5 b . A remarkable small ichthyosaur from the
. 1989c. Leptopterygius tenuirostris and other long- Upper Triassic of British Columbia, representing
snouted ichthyosaurs from the English Lower a new genus and species. Can. Jour. Earth Sci.
Lias. Palaeontology 32(3)409—27. 32:292-303.
Eurhinosaurus (Reptilia Ichthyosauria) does have a Hucnc, 1 9 2 2 (Reptilia: Ichthyosauria). Can. Jour.
tailbend. Can. jour. Earth Sci. 2 7 ( n ) : i 5 4 i — 4 5 . Earth Sci. 3 3 : 4 3 9 — 4 3 .
McGowan, C, and A. C. Milner. 1 9 9 9 . A new ichthyopterygian forefins. four. Vert. Paleo. 19(1)28—
20(2)295—
McNab, B. K. 1 9 8 3 . Energetics, body size, and the
. 2 0 0 0 b . Rulers of the Jurassic seas: T h e reign of
limits to endothermy. Jour. Zoo/. London 199:1—29.
Plcsiosauroidea). Bull. Inst. Roy. Sci. Nat. Belg. Nicholls, E. L., D. B. Brinkman, and J. M. Callaway.
70:161—78. 1999. New material of Phalarodon (Reptilia:
Naish, D. 1998. Did ichthyosaurs fly? Dinosaur World Ichthyosauria) f r o m the Triassic of British
4:27-29. Columbia and its bearing on the
Naish, D, L. F. Noe, and D. M. Martill. 2 0 0 1 . Giant interrelationships of mixosaurs. Palaeontographica
reptile from Bedfordshire. Animals io(2):6i—63. Noe, L . 1 9 9 9 . T h e Callovian pliosaurs o f the Oxford
Nicholls, E. L. 1 9 7 6 . T h e oldest known North Clay —evidence and implications for the
American occurrence o f the Plesiosauria consumption of marine invertebrates. In E. Hoch
(Reptilia: Sauropterygia) f r o m the Liassic (Lower and A. K. Brantsen, eds., Secondary Adaptation to Life
Jurassic) Fcrnic Group Alberta, Canada. Can. Jour. in Water, pp. 38—40. University o f Copenhagen
Earth Sci. 13:185-88. Gcologisk Museum.
. 1 9 8 8 . T h e first record of the mosasaur . 2 0 0 1 . A taxonomic and functional study of the
Hainosaurus (Reptilia: Laccrtilia) from North Callovian (middle Jurassic) Pliosauroidca
America. Can. Jour. Earth Sci. 25:1564—70. (Reptilia, Sauropterygia). Ph.D. diss., University
Nicholls, E. L., and D. B. Brmkman. 1 9 9 3 . A new of Derby.
specimen of Utatsusaurus (Reptilia: Norman, D. 1994. Prehistoric Life. Macmillan.
Ichthyosauridae) from the Lower T r i a s s i c Sulphur Novacck, M . J. 1996. Dinosaurs of the Flaming Cliffs.
from flipper geometry: Parallels among birds, Palaeont. Soc. London 111:1—12.
bats, and airplanes. Jour. Vert. Paleo. 22 (suppl. to . 1 8 7 7 . On the rank and the affinities in the
3> 3A. 9
reptilian class of the Mosasauridae, Gervais.
OrndorfT, R. L., R. W. Wieder, and H. F. Filkorn. Quart. Jour. Geol. Soc. London 33:682—715.
. 1 8 4 0 . Note on the dislocation of the tail at a Pearce, F. 2 0 0 2 . Oldest fossilized vomit pile
. 1 8 4 2 . Observations on the teeth of the Ichthyosaurus (communis?). Ann. Mag. Nat. Hist. 17:44—
Soc. London 3:24—28. Perkins, S. 2002. Sea dragons. Science News i62(8):i22—
Association for the Advancement of Science. John Murray. Persson, P. O. 1 9 8 2 . Elasmosaurid skull from the
Lower Cretaceous of Queensland (Reptilia: Denmark. In E. Hoch and A. K. Brantsen, eds.,
Sauropterygia). M e m . Q l d . M u s . 2 0 : 6 4 7 — 5 5 . Secondary Adaptation to Life in Water, p. 70. University
Pitman, R. L, D. M. Palacios, P. L. R. Brennan, B. J. of Copenhagen Gcologisk Museum.
Brcnnan, K. C. Balcomb, and T. Miyashita. 199c). Reeves, R. R., B. S. Stewart, P. J. Clapham, and J. A.
Sightings and possible identity of a bottlenose Powell. 2002. National Audubon Society Guide to Marine
45:1—104.
Raup, D. M., and J. J. Scpkoski. 1 9 8 2 . Mass Geol.
215:1501-3.
Sauropterygia) f r o m the Upper Muschelkalk
Raup, D. M., and S. M . Stanley. 1971. Principles of Ricppcl. O., and H. Hagcdorn. 1 9 9 7 . Paleogeography
Mass extinctions in the fossil record. Science Nicholls, eds., Ancient Marine Reptiles, pp. 121—44.
concerning plesiosaur locomotion. N. Jb. Ceol. Rowe, M . P. 2 0 0 0 . Inferring the retinal anatomy and
Palaeont. 164:193—94. visual capacities of extinct vertebrates. Palaeontologia
that got away? Aust. Nat. Hist. 19:408—13. University of Chicago Press.
. 1988. W h o should pay for Australia's past? Aust. . 1992. Scenes from Deep Time. University o f
Nat. Hist. 22(8)368-70. Chicago Press.
. 1990. Return of the great sea monsters. Aust. . 1997. Ceorgcs Cuvier, Fossil Bones, and Geological
. 1966. Vertebrate Paleontology. University of Chicago . 1975a. A new species of Clobidens from South
Press. Dakota, and a review of globidentine mosasaurs.
. 1968. The Procession of Life. World. Fieldiana Ceol. 33(13)235—56.
skeleton of the giant plesiosaur Kronosaurus. Interior of North America, pp. 119—36. Geological
Breviora 112:1—15. Association of Canada.
Roper, C. F. E., and K. J. Boss. 1 9 8 2 . T h e giant squid. . 1989. An Odyssey in Time: The Dinosaurs of North
Nothosauria) from the middle Triassic of Monte Scanlon, J. D., M. S. Y. Lee. M. W. Caldwell, and
San Giorgio (Switzerland), with the description
R. Shine. 1 9 9 9 . T h e palcoccology of the primitive
of a new species. Phil. Trans. Roy. Soc. London B
snake Pachyraehis. Llistorical Biology 13:127—52.
325:561—670.
Schwimmcr, D. R., J. D. Stewart, and G. D. Williams.
. 1992. Cymbospondylus (Shastasauridae:
1997. Scavenging by sharks of the genus Sijualieorax
Ichthyosauria) from the middle Triassic of
in the late Cretaceous of North America. Palaios
Spitsbergen: Filling a palcobiogcographic gap.
12:71-83.
Jour. Paleo. 66(2)332—37.
Scott, W. B., and S. N. Tibbo. 1 9 6 8 . Food and feeding
. 1997. T h e paleobiogcography of Shastasaurus. In
habits of the swordfish. Xiphiasgladias, in the
J. M . Callaway and E. L. Nicholls, eds., Ancient
western North Atlantic, four. Lisb. Res. Bd. Can.
Marine Reptiles, pp. 17—43. Academic Press.
25(5>9°?-'9-
. 2 0 0 0 . Ichthyosauria: Their diversity,
Seeley, H. G. 1874. On Muraenosaurus leedsi, a
distribution, and phylogeny. Palaontol. Z. 74(1—
plesiosaurian from the Oxford Clay. Q. Jour. Geol.
2)1-35.
Soc. London 30:197—208.
Sander, P. M., and H. Buchcr. 1 9 9 0 . On the presence
. 1877a. On Mauisaurus gardneri, Seeley, an
of Mixosaurus (Ichthyopterygia: Reptilia) in the
elasmosaurian from the base of the Gault of
middle Triassic of Nevada. Jour. Paleo. 64(i):i6i—
64.
Folkestone. Q. Jour. Geol. Soc. London 33:541—46.
Mh. 10:631—40.
Clay of St. Ncots. Q. Jour. Geol. Soc. London 30:716—
. 1 9 9 2 . Ontogenetic changes in mosasaur bone and Conservation of Sea Turtles, pp. 591—92.
microstructure. your. Vert. Paleo. I2(suppl. to 3):5iA. Smithsonian Institution Press.
. 1 9 9 7 . Ecological implications of mosasaur bone Spotila, J. R., M. P. O'Connor, and F. P. Paladino.
microstructure. In J. M. Callaway and E. L. 1997. Thermal biology. In P. L. Lutz and J. L.
Nicholls, eds., Ancient Marine Reptiles, pp. 333—54. Musick, eds.. The Biology of Sea Turtles, pp. 297—314.
Academic Press. C R C Press.
Sheldon, M. A., and G. L. Bell. 1 9 9 9 . Stanley, S. M. 1984a. Marine mass extinctions: A
Pacdomorphosis in Mosasauridae (Squamata): dominant role for temperature. In M. H. Nitccki,
Evidence from fossil bone microstructure. ed., Extinctions, pp. 69—117. University of Chicago
Paludicola 2(2)190—205. Press.
Sheldon, M. A., G. L. Bell, and J. P. Lamb. 1 9 9 6 . . 1984b. Mass extinctions in the ocean. Scientific
Simpson, G. G. 1953. Life of the Past. Yale University Deer River, Alherta, Canada. Lawrence, Kans.: n.p.
Press. 1985 reprint, NeWcst Press.
. 1 9 2 2 . Explorations of the Permian of Texas and Storrs, G. W, and M. A. Taylor. 1996. Cranial
the chalk of Kansas, 1918. Trans. Kansas Acad. Sci. anatomy of a new plesiosaur genus from the
30(1)119—20. lowermost Lias ( R h a e t i a n / H e t t a n g i a n ) of Street,
Stewart, J. D., and G. L. Bell. 1989. T h e earliest Somerset, England. Jour. Vert. Paleo. 16:403—20.
reputed North American mosasaur records are Storrs, G. W, M. S. Arkhangelsk!!, and V M.
not mosasaurs.your Vert. Paleo. 9:j9A (abstract). Efimov. 2000. Mesozoic marine reptiles of Russia
. 1994. North America's oldest mosasaurs are and other former Soviet republics. In M. J.
teleosts. Contrib. Sci. Nat. Hist. Mus. L A . County
Benton, M. A. Shiskin, D. M. Unwin, and E. N.
441:1-9. Kurichkin, eds., The Age of Dinosaurs in Russia and
Storrs, G. W. 1984. Elasmosaurus platyurus and a page Mongolia, pp. 187—210. Cambridge University Press.
from the Cope-Marsh war. Discovery 17(2)25—27.
Stukcly, W. 1719. An account of the impression of an
. 1991. Anatomy and relationships of Corosaurus
almost entire skeleton of a large animal in a very
aleovensis (Diapsida: Sauropterygia) and the
hard stone from Nottinghamshire. Phil. Trans. Roy.
clarification of the genus Plesiosaurus. In J. M . . 1930b. On fossil reptilia from Sokoto province.
Callaway and E. L. Nicholls, eds., Ancient Marine Biol. Ceol. Surv. Nigeria Bull. 1:42—48.
Reptiles, pp. 145—90. Academic Press. . 1930c. Preliminary account of a new genus and
. 1 9 9 9 . An examination of plesiosauria species of plesiosaur. Ann. Mag. Nat. Hist.
. 1973. Fossil Reptiles and Amphibians. British Museum. . 1 9 9 3 . Stomach bones for feeding or buoyancy?
Tarlo, L. B. 1958. The scapula of Pliosaurus macromerus, T h e occurrence and function of gastroliths in
Phillips. Palaeontology 1:193—99. marine tetrapods. Phi! Trans. Roy. Soc. London B
. 1959b. Stretosaurus gen. nov., a giant pliosaur Institution and its contribution to vertebrate
. i 9 6 0 . A review of the Upper Jurassic pliosaurs. . 1995. T h e sea monster of Dorset. Natural History
104(10)69.
Bull. Br. Mus. (Nat. Hist). Geol. 4 (5):i47-8 . 9
. 1 9 8 7 b . A reinterprctation of ichthyosaur
Taylor, M. A., D. B. Norman, and A. R. I.
swimming and buoyancy. Palaeontology 30(33:551—35.
Cruickshank. 1 9 9 3 . Remains of an ornithischian
. 1 9 8 7 c . Reptiles that took on the sea. N e w dinosaur in a pliosaur from the Kimmeridgian of
Scientist 116(15883:46—50. England. Palaeontology 36(2)357—60.
. 1989a. T h e other dinosaurs. New Scientist Tchernov, E., O. Ricppel, H. Zahcr, M. J. Polcyn, and
121(1655)65. L. J. Jacobs. 2000. A fossil snake with limbs. Science
. 1 9 9 2 . Functional anatomy of the head of the Thompson, K. S., and D. E. Simanck. 1 9 7 7 . Body
large aquatic predator Rhomaleosaurus zetlandicus form and locomotion in sharks. American Zoologist
bitten by a pliosaur. Modern Geology 18:489—501. Wade, M . 1984. Platypterygius australis, an Australian
Thurmond, J. T. 1 9 6 8 . A new polycotylid plesiosaur Cretaceous ichthyosaur. l.ethaia 17:99—113.
from the Lake Waco Formation (Cenomanian) of . 1 9 9 0 . A review of the Australian Cretaceous
Texas. Jour. Paleo. 42:1289—96. longipinnate ichthyosaur Platypterygius
. 1969. Notes on mosasaurs from Texas. Texas (Ichthyosauria, Ichthyopterygia). Mem. Old. Mus.
Jour. Sci. 2i(i):69—79. 28:115-37.
T h u r m o n d , J. T , and D. E. Jones. 1981. Fossil Vertebrates Wagner, A. 1 8 5 3 . Die characteristic eine ncuen art von
of Alabama. University of Alabama Press. Ichthyosaurus aus den Iithograpfuschen schicfcrn
Tibbo, S. N., L. R. Day, and W. F. Doucet. 1 9 6 1 . T h e und eines zahnes von Polyptyehodon aus dem
swordfish (Xiphiasgladius L.): Its life history and griindsandsteine von Kellhcim. Bull, der koniglische
economic importance in the Northwest Atlantic. Akademie der Wissenschaft, Gelehrte Anzeigen 6:661—70.
Bull. Fish. Res. Bd. Can. 130:1—47. Ward, P. D. 1992. On Methuselah's Trail. Freeman.
Tichy, G. 1 9 9 5 . Ein fruhcr, durophager Ichthyosaurier . 1997. The Call of Distant Mammoths. Copernicus.
(Omphalosauridae) aus der Mitteltrias der Alpen. . 1998. Time Machine: Scientific Explorations in Deep
6:167-78. 917.
Tokaryk, T. T. 1 9 9 3 . A plioplatecarpine mosasaur Watson, L. 1981. Sea Guide to Whales of the World.
limpets responsible for the perforations in the Welles, S. P. 1 9 3 9 . Plesiosaur from the Upper
ammonite Placenterias? Palaeogeography, Cretaceous of the San Joaquin Valley. Bull. Geol.
51(2)1424-25.
Wilford, J. N. 1986. The Riddle of the Dinosaurs. Knopf.
queenslandicus Longman. Occasional Papers Boston Soc. on the Cretaceous paleogeography of North
. 1940. Holotype of Plesiosaurus longirostris Blake System in the Western Interior of North America, pp. I—
and the classification of the plesiosaurs. Jour. Paleo. 2 0 . Geological Society of Canada.
Ann. Mag. Nat. Hist. 9:742—44. . 1 8 8 9 . A new plesiosaur from the Niobrara
Cretaceous of Kansas. Trans. Kansas Acad. Sci. Winchester, S. 2001. The Map that Changed the World:
. 1 8 9 7 . Range and distribution of the mosasaurs (Squamata, Osasauridae) from the eastern United
with remarks on synonymy. Kansas Univ. Quart.
States, your Vert. Paleo. 8(5)545—45.
4 ( 4 > 7 7 - 8 5 -
Wright, K. R., and S. W. Shannon. 1988. Selmasaurus
. 1914. Water Reptiles of the Past and Present. design, fabrication, engineering and installation of
Wilson, E. O. 1992. The Diversity of Life. Harvard plesiosaur).your Vert. Paleo. I5(suppl. to 5)161 A.
Wiman, C. J. 1 9 2 0 . Some reptiles from the Niobrara Egypt and in Africa in general. Bull. Inst. Egypte
asiaticus, 94
1J7
Dawson, J . W , 2 8 , 4 2 , 165
fossils collected by, 2 0 7 — 2 0 8 , 2 1 0 , 2 1 1 — 2 1 3 , 2 1 9
DcBraga, Michael, 2 0 0 , 2 0 1
mosasaur names, 2 0 8 , 2 0 9 , 2 1 0
Decompression sickness. See Bends
o n mosasaurs, 2 1 0 , 2 1 9 , 2 4 5
Deeming, D. G, 4 9 , 7 3 — 7 4
plesiosaur species identified, 1 2 9 , 1 3 0
Dcinonychus, 46, 255
rivalry with Marsh, 1 2 7 — 1 2 8 , 2 1 0 , 2 1 1 — 2 1 3 , ' 9 2
Creationists, 6 3 — 6 4
Desmond, Adrian J., 4 6
Diapsids, 2 8 — 2 9 , 2 0 4
Crcislcr, Ben, 1 6 8 — 1 6 9 , ' 8 6 , 1 8 8
Dinilysia, 247
Cretaceous era, 1 1 — 1 2
Cretoxyrhina mantelli (lamnid shark), 241—242
Dinosaurs
marine, 4 4 , 5 8 , 1 0 4 diets, 4 2
flippers, 14 Elephants, 2 4 , i 8 2 n
hunting behavior, 2 ( 9 Enaliosaurs, 166
leaping, 101 England
locomotion, 4 1 , 103 dinosaur fossils, 1 9 — 2 0
similarities to ichthyosaurs, 11, 1 4 — 1 5 , 1 6 , 4 1 , 5 0 — 5 2 , fossil sites, 9
61, 62(illus.), 7 m , 100—10m ichthyosaur fossils, 6 5 — 6 6 , 6 7 — 7 3 , 8 1 — 8 2 , 1 0 5 — 1 0 7
Dunklcosteus, 2 Enzymes, 5 6 — 5 7
Great white sharks (Carcbarodon earebarias), 59, 79, 216, Holzmaden, Germany, 9 , 7 4 — 7 5 , 8 1 , 8 6
Home, Sir Everard, 6 7 — 6 8
o f ichthyosaurs, 8 0 , 9 9 Macroplerygius, 96
of plesiosaurs, 1 2 0 , 1 4 3 , 1 5 2 — 1 5 4 Mollesaurus, 95
Hylaeosaurus, 25 Quamcbtbyosaurus, 96
in Argentina, 95 Svalbardosaurus, 80
fetuses, 7 2 — 7 4 , 81 Wimanius, 86
in Germany, 7 4 — 7 5 , 8 1 , 8 6 , 9 6 — 9 7 , 1 1 4 Ichthyosaurs
in New Zealand, 2 2 7 aquatic adaptation, 4 3 , 6 4 — 6 5
Iehthyosaur genera body plans, 14—15, 4 4 , 4 8 , 50—52, 61—63, 62(illus.
Aegirosaurus, 96—97 64, 79—80, 104, in
Chacaisaurus, 95 colors, 9 9 — 1 0 0
Isfjordosaurus, 84 eyes, 7 7 , 9 6 , 9 9 , 1 0 5 , 1 0 9 — 1 1 1
leaping, 1 0 0 — 1 0 2 Israel
Kronosaurus, 175—179, 180, 185, 192, 193 Lingham-Soliar, Thcagartcn, 15, 16, 7 9 , 115—116, 1 4 1 —
attacks on elasmosaurs, 1 3 2 142, 1 4 5 — 1 4 7 , 148, 196, 197, 198, 200—201, 214,
K-T extinction. See Asteroid impact; Extinctions Linnaeus, Carolus (Carl von Linne), 3, 1 7 — 1 8 , 19
Liodon, 210, 250
Leatherback turtles (Dermochelys), 32, 33, 53(illus.), 53— See also Leiodon
kempii, 34 teeth, 4 3 , 1 7 9 , 1 8 0 , 1 8 4
Locomotion posthumus, 96
aquatic, 4 8 trigonus, 96
methods, 1 3 6 — 1 3 7 Mammals
175
marine, 5 2 — 5 3 , 5 4
sauroptcrygians, 3 4 — 3 6 cctothcrmic, 2 7 — 2 8 , 2 9 , 4 5 , 52
Pliosaurs of mosasaurs, 2 3 8 — 2 3 9
McHenry, Colin, 5 1 — 5 2 , 1 2 0 , 1 7 7 , 186, 191 Monitor lizards (Megalama prisca), 58—59, 239
3 0 2 S' £ A 'D %A g O T V 5
fossils found in, 1 9 8 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
embryos, 2 1 7 flippers, 2 0 4
first discoveries, 7 , 2 4 , 1 9 5 — 1 9 8 , 251 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
213, 215, 2 1 6 , 2 1 8 — 2 1 9 , 2 4 1 — 2 4 2 , 2 5 0 latest species, 2 4 4 , 2 5 7
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
J NT) EX 303
Mosasaurus, continued ichthyosaur fossils, 2 2 7
boffmanni, 7, i96(illus.), 197—198, 216, 220, 226, 229, mosasaur fossils, 2 0 3 , 2 2 7 — 2 2 8 , 2 4 4
2 3 0 — 2 3 1 , 251 plesiosaur fossils, 1 0 , 134—135
horridus, 216 pliosaur fossils, 9 — 1 0
lemonnieri, 221 Nicholls, Elizabeth L., 55, 8 9 - 9 0 , 9 3 , 9 6 , 2 2 o n , 2 3 0 ,
maxitniliani, 198
Necks nevadanus, 87
fat, 53
184
175
Dolichorhynchops, 130
name, 1 9 , 6 6 , 1 1 8 , 1 2 3 — 1 2 4
Hydraltnosaurus, 130
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
Hydrotherosaurus, 122
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
Kaiwbekea, 134—135
possible ancestors, 36—37, 1 2 0
Kimmerosaurus, 136
reproduction, 1 4 7 — 1 4 8 , 1 4 9 — 1 5 1 , 1 5 2
130, 159—160
Libonectes,
ribs, 8 6 , 1 1 9
Muraenosaurus, 121, I22(illus.), 132, 135, 184
skin,161
130
Polycotylus,
skulls, 4 3 , i33(illus.)
130
Styxosaurus,
smallest, 150
160—161
Tkalassiodracon,
tail fins, 15, 138, 1 4 2
Tbalassiosaurus, 130
taxonomy, 1 2 9 — 1 3 0 , 1 3 2 — 1 3 3 , 1 6 0
Thalassomedon, 2—3, 129, 130—132, i3i(illus.) teeth, 4 3 , 118, 1 3 4 , 135, 136, 1 6 6 , 1 8 4
Thalassonomosaurus, 130
time period, 4 4
132, 135—136
Trideidus,
unanswered questions about, 151—153
Tuarangisaurus, 10
vertebrae, 1 2 0 , 135, 1 6 0
132, 178—179
Woolungasaurus,
See also Elasmosaurs; Marine reptiles; Pliosaurs
See also Elasmosaurus; Plesiosaurus
Plesiosaurus, 118, 124
dolichodeirus, 4, 4(illus.\ 19, 69, I2j(illus.), 125—124, 5c? a/so Liopleurodon; Pliosaurus
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
in New Zealand, 9 — 1 0 reproduction, 150—151
tuckeri, 214
Polycotylus, 192
Plurtdens walken, 114—11%
Polyptychodon, 192, 193
Polycotylidac, 189, 192
Rhomaleosaurus, 169, i7o(illus.), 171—172, 174, 192
Polycotylus, 192
Sitnolcstes, 187, 188, 192
latipinnis, 130
Stretosaurus, 185
Polyptychodon, 192, 193 definition, 2 7
Porbeagle sharks (Lamna nasus), 62(illus.), 79 ectothermic nature, 2 7 — 2 8 , 2 9 , 4 5 , 52
Powell, James L., 2 5 4 eggs, 2 7
Proaigialosaurus hueni, 200, 202 energy needs, 54—55
Proganockilys quenstedi, 31, 32 evolution of, 2 9 , 30
Prognathodon, 198—199, 219, 225, 229 lack o f stamina, 4 7
currii, 226—227 return to aquatic life, 30
giganteus, 221 ribs, 6 3 n
overtoni, 227 ruling, 4 4
saturator, 221—222, 222(illus.) Sphenodontidac, 84—85
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 ,
Romer, Alfred Sherwood, 6 3 , 1 7 5 , 2 4 5
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
Roux, Erica, 2 3 6
of marine reptiles, 1 4 , 2 9 — 3 0 , 4 8 — 4 9 , 58
Rudwick, Martin J . S., 2 1 , 2 2 , 2 4 , 7 0
o f mosasaurs, 1 9 9 , 2 1 7 , 2 1 8
Ruling reptiles, 4 4
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
Russell, A. P., 55
of pliosaurs, 150—151
Russell, Dale A., 1 9 8 , 2 0 0 , 2 0 5 , 2 0 7 , 2 1 0 , 2 1 3 , 2 1 4 , 2 1 5 ,
viviparous, 2 9 — 3 0 , 4 3 , 4 8 — 4 9 , 5 8 , 7 2 — 7 4 , 9 7 — 9 8 ,
217, 219, 221, 239, 2 4 2 , 2 4 4
99, 217—218
Russia, fossils found in, 181
Reptiles
Ruta, Marcello, 2 4 9
anapsids and diapsids, 2 8 — 2 9 , 5 8 n
endangered, 34 altispinus, 89
flatback, 34 earinthiaeus, 89
skulls, 2 4 8 — 2 4 g Stupendemysgeographicus, 32
of snakes, 2 4 9 Tuataras, 2 7 , 8 4
of whales, 1 7 6 Tuna