R. McNeill Alexander-Dynamics of Dinosaurs and Other Extinct Giants-Columbia University Press (1989) PDF

DYNAMICS OF
DINOSAURS
& OTHER EXTINCT GIANTS
R. MCNEILL A L E X A N D E R
R. M C N E I L L A L E X A N D E R
Dynamics of Dinosaurs
and Other Extinct Giants
Columbia University Press

NEW YORK
C O L U M B I A U N I V E R S I T Y PRESS
N e w York G u i l d f o r d , Surrey
C o p y r i g h t © 1989 C o l u m b i a U n i v e r s i t y Press
All rights reserved
LIBRARY OF C O N G R E S S
Library of C o n g r e s s C a t a l o g i n g - i n - P u b l i c a t i o n D a t a
A l e x a n d e r , R. M c N e i l l .
D y n a m i c s of d i n o s a u r s a n d o t h e r e x t i n c t g i a n t s / R. M c N e i l l A l e x a n d e r .
p. cm.
Includes bibliographies and index.
ISBN 0 - 2 3 1 - 0 6 6 6 6 - X
1. D i n o s a u r s — L o c o m o t i o n .
2. Extinct a n i m a l s — L o c o m o t i o n .
3. Animal mechanics.
I. T i t l e .
Q E 8 6 2 . D 5 A 3 3 1989
567.9T—dcl9 88-20373
CIP
P r i n t e d in t h e U n i t e d S t a t e s of A m e r i c a
C a s e b o u n d e d i t i o n s o f C o l u m b i a U n i v e r s i t y Press b o o k s are S m y t h - s e w n
a n d p r i n t e d o n p e r m a n e n t a n d d u r a b l e acid-free paper
CONTENTS
PREFACE VU
I. Introducing the Dinosaurs 1

II. Weighing the Dinosaurs 16
III. Dinosaur Footprints 27
iv. Dinosaur Strength 44
v. Dinosaur Necks and Tails 60
vi. Fighting and Singing Dinosaurs 73
VII. Hot-Blooded Dinosaurs? 90
vin. Flying Reptiles 106
ix. Marine Reptiles 122
x. Death of the Giant Reptiles 141
xi. Giant Birds 150
XII. Giant M a m m a l s 159
xm. Epilogue 162

INDEX 165
PREFACE
T H I S is a book about big, extinct animals. Most of it is about di-

nosaurs, including the biggest land animals that have ever lived.
There is a chapter about the huge reptiles that lived in the sea in the
time of the dinosaurs and one about the biggest of the flying reptiles
that lived at the same t i m e . There are also short chapters about gigan-
tic, extinct birds and m a m m a l s . Some very big reptiles lived before the
dinosaurs but I have written nothing about them, because all of t h e m
were m u c h smaller than the biggest dinosaurs and because fewer peo-
ple are interested in t h e m .
There are many other books about dinosaurs but none, I think, like
this one. I have used the m e t h o d s of physics and engineering to try to
discover how extinct animals could have lived and moved. I like to
think about animals in the kinds of ways that engineers think about
machines and vehicles. T h a t seems the best way of finding out how
dinosaurs could have worked.
You do not need to k n o w m u c h science to understand this book. I
have tried always to start from basics, and to keep the arguments sim-
ple. However, if you do already k n o w a lot of science, please do not be
put off by the simple explanations, because I t h i n k you will find ideas
here that you have not read elsewhere. T h i s book is m e a n t for everyone
who is interested in dinosaurs: scientists and nonscientists, school-
children and professors.
I
Introducing the Dinosaurs
T tells what dinosaur fossils are like and how they are
HIS C H A P T E R
formed and introduces some of the best-known dinosaurs. It is an

introductory chapter intended m a i n l y for readers w h o have little pre-
vious knowledge of these remarkable reptiles. Other may prefer to skip
quickly through the chapter or to go directly to chapter 2.
Figure 1.1 shows a dinosaur 13 meters (43 feet) tall, well over twice
the height of a full-grown (5.5 meter) giraffe. It is far bigger than any
modern land animal and its skeleton (now in Berlin) is one of the world's
most impressive m u s e u m exhibits.
This dinosaur, Brachiosaurus, is the largest k n o w n with a reason-
ably complete skeleton, but may not be the largest that ever lived. Pos-
sible rivals such as Supersaurus and Ultrasaurus are k n o w n by only a
few bones each, and scientists are still arguing about the sizes of the
complete dinosaurs.
This book is about gigantic animals, so most of the dinosaurs in it
are very big ones, but figure 1.2 shows that there were smaller ones
too. The Compsognathus was about the size of a hen and the Psitta-
cosaurus was smaller than a pigeon, but both of t h e m were juveniles.
The Compsognathus skeleton (including the tail) is 0.8 meters long,
which is not too m u c h less than the 1.4 m e t e r s of another specimen
that is probably adult. T h e Psittacosaurus, however, belongs to a spe-
cies that grew to an adult length of 2 meters, w h i c h is still small for
a dinosaur.
Figures 1.1 and 1.2 show dinosaurs as they are thought to have looked
in life, but the actual specimens are mere skeletons. Figure 1.3 shows
one of the fossils of Psittacosaurus (an adult). Some of the leg and tail
bones are missing but have been drawn in, as they are thought to have
looked, with dotted lines. The skeleton has collapsed or been squashed
after death, making ribs stick out at odd angles, but the bones have
2 INTRODUCING THE DINOSAURS
stayed together. Many other dinosaur finds have been of scattered or

jumbled bones.
Many big dinosaur skeletons have been separated bone by bone from
the rock and reassembled as complete skeletons in natural postures.
T h e Psittacosaurus skeleton (figure 1.3) was given m u c h less drastic
treatment. Only enough rock was removed to reveal all the bones, which
were left attached to the remaining block of stone. The skeleton in
natural posture (figure 1.4) was not actually constructed but the draw-
ing was built up from drawings of the individual bones. Making draw-
ings like this is an important stage in reconstructing the appearance of
extinct animals. Adding flesh and skin to m a k e drawings like figure
1.2 depends largely on knowledge of the soft a n a t o m y of living ani-
mals, but also requires a good deal of guesswork.
Fossils get preserved because the earth's surface is continually
changing. Massive earth m o v e m e n t s crinkle it up into chains of moun-
tains, which get worn down again by the processes of erosion. The
Rockies, the Andes and the Alps are all more recent than the dino-
saurs, and older m o u n t a i n s , such as the Appalachians, are less im-
pressive because there has been more t i m e for t h e m to be eroded. The
next few paragraphs explain how m o u n t a i n building and erosion pre-
serve fossils and m a k e t h e m possible to find.
Exposed surfaces of rocks get heated by the sun during the day and
F I G U R E 1.1. Brachiosaurus w i t h a h o u s e a n d a n a d u l t giraffe d r a w n t o t h e

s a m e scale. T h e d i n o s a u r , w h i c h is 13 m e t e r s tall, is d r a w n from a m o d e l sold
by the Natural History M u s e u m , London.
INTRODUCING THE DINOSAURS 3
FIGURE 1.2. J u v e n i l e Compsognathus ( b e h i n d ) a n d Psittacosaurus (right), w i t h

a domestic pigeon. T h e dinosaur skeletons on w h i c h the drawings arc based
are a m o n g t h e s m a l l e s t k n o w n . D r a w n b y M a t t h e w H y m a n . F r o m C o o m b s 1980.
R e p r i n t e d by p e r m i s s i o n . C o p y r i g h t © 1980 M a c m i l l a n M a g a z i n e s Ltd.
cool during the night. This makes t h e m expand and contract, breaking
fragments off. Water seeps into cracks in rocks and freezes in winter,
swelling and splitting the rock. Sand grains carried along by streams
scour away the rock of the stream bed. These processes break rocks
down into sand grains and m u d particles which are washed away by
rainwater and streams, which carry t h e m down toward the sea. T h e
sand and mud settle out where the water runs more slowly, on flooded
plains or on sand banks or m u d flats at a river's m o u t h . Carcasses of
animals that die in these places may get buried in the sand or mud
(figure 1.5a-c). Carcasses also get buried in sand dunes: that is what
happened to the Psittacosaurus, w h i c h seem to have lived in a dry,
sandy area. Yet other carcasses get buried in the calcium carbonate
which, in certain circumstances, precipitates out in the sea. I do not
know of any dinosaur fossils that got preserved like that but m a n y of
the fossils of marine reptiles, described in chapter 9, are in limestones
formed from calcium carbonate precipitates. T h e flesh of carcasses rots
away but buried skeletons may be preserved. Skeletons that do not get
buried are generally destroyed, broken down by the same weathering
processes as erode rocks.
As time passes and more sediment accumulates, skeletons get buried
deeper and deeper (figure 1.5d). T h e sediments consolidate into rock:
sand becomes sandstone, m u d becomes shale and calcium carbonate

precipitates become limestone. The fossil may come to lie in solid rock,
deep below the ground. If it stayed there it would never be found unless
perhaps by mining or quarrying. However, earth m o v e m e n t s crinkling
the earth's surface m a y raise it up in a m o u n t a i n (figure 1.5e) and ero-
sion, wearing the m o u n t a i n away, m a y expose it (figure 1.5f). If those
things happen it m a y be found, but some of its bones may get broken
or worn away first. The Psittacosaurus in figure 1.3 had lost parts of
its feet by erosion, before it was found.
Usually only skeletons survive as fossils, but there are sometimes
traces of other parts. If a carcass gets buried before the soft parts decay,
the sediment will mould itself to the skin surface and in rare cases
this impression of the skin may survive. Impressions of skin have been
found with several fossils of duck-billed dinosaurs (dinosaurs like the
one shown in figure 1.11). They show that the skin was scaly like tor-
toise or lizard skin. Fossil stomach c o n t e n t s are s o m e t i m e s found: for
example, one of the Compsognathus fossils has a lizard skeleton inside
it. There are also some fossil dinosaur eggshells, which resemble the
eggshells of birds and modern reptiles. T h e most famous belong to Pro-
toceratops (a close relative of Psittacosaurus) as fossil embryos found
with t h e m show. Adult Protoceratops were 2 meters long and the eggs
measure 1 0 - 2 0 centimeters. They seem to have been laid in hollows
in the ground in clutches of thirty or more. Finally, there are a lot of
dinosaur footprints: they will be described in chapter 3.
Figure 1.5 shows two dinosaur skeletons getting buried, one after the
other. By the time the small one died (figure 1.5c) the big one was
F I G U R E 1.3. A Psittacosaurus s k e l e t o n a s i t w a s f o u n d , with enough rock

cleared a w a y t o s h o w t h e b o n e s . F r o m O s b o r n 1924.
F I G U R E 1.4. A r e c o n s t r u c t i o n of t h e s k e l e t o n of Psittacosaurus, b a s e d on t h e
s k e l e t o n s h o w n i n f i g u r e 1.3. T h e o v e r a l l l e n g t h o f t h e s k e l e t o n w a s 1.4 m e t e r s .
From O s b o r n 1924.
already buried, so the small one finished up in a higher layer of rock.

It would be obvious to a paleontologist w h o found both fossils that the
big one was the earlier of the two.
The relative ages of fossils are less obvious w h e n they are found far
apart, possibly in different continents, but they can be worked out by
an extension of the same principle. Layers of rock of the same age all
over the world can be matched up by the similarity of their fossils.
Nineteenth-century geologists discovered this and used it to divide time
up into four eras: Precambrian (the oldest), Palaeozoic, Mesozoic, and
Cenozoic (table 1.1). Fossils are rare in Precambrian rocks but are plen-
tiful in rocks from the later eras. All the dinosaurs lived in the Me-
sozoic era. Each era is subdivided into periods. T h e Mesozoic consists
of the Triassic, Jurassic, and Cretaceous periods, w i t h dinosaurs living
in all three.
The periods were given n a m e s before anyone could tell how long
each of t h e m had lasted, but the discovery of radioactivity made dating
possible. Radioactive isotopes break down into other isotopes, each at
its own characteristic rate. Some break down exceedingly slowly, over
periods of hundred of millions of years. W h e n they are found in rocks
it is sometimes possible to calculate the rock's age from the proportion
of original isotope to breakdown products. Such m e a s u r e m e n t s tell us
that the Mesozoic era began about 230 million years ago and ended 65
million years ago. T h e dates are s h o w n in table 1.1.
Each dinosaur species lived for only a small part of the Mesozoic.
For example, Compsognathus longipes (figure 1.2) lived late in the Jur-
assic period, about 140 million years ago, and Psittacosaurus mongo-
liensis lived in the Cretaceous period, about 90 million years ago. Ta-
ble 1.2 shows when these and other dinosaurs lived.
I gave each species its full n a m e in that paragraph, because I wanted
to emphasize that I was talking about single species. Every animal spe-
cies, w h e t h e r living or fossil, is given two names, which usually have
meanings derived from Latin or Greek. Psittacosaurus means "parrot-
lizard" (notice its parrot-like beak) and mongoliensis means "mongo-
l i a n " (telling where the fossils were found), so the two n a m e s together
mean "mongolian parrot-lizard." Similarly Compsognathus longipes
means "long-footed pretty jaw." The first name in each case is the name
of the genus, which may include several closely related species (for
example lions Panthera leo and tigers Panthera tigris are both m e m -
bers of the genus Panthera). If there are several species, the second
n a m e tells us which is m e a n t . It so happens that only one species of
Psittacosaurus is known, and one of Compsognathus, so it will gen-
erally be unnecessary to use the second names until more species are
discovered. However, several species of Brachiosaurus have been rec-
ognized, including Brachiosaurus altithorax from Colorado and Brach-
iosaurus brancai from Tanzania. Even in cases like this it is unnec-
essary to use the second n a m e if it does not m a t t e r which of the closely
similar species is being referred to. Most of the things that I might
F I G U R E 1.5. D i a g r a m s s h o w i n g h o w d i n o s a u r s h a v e b e e n f o s s d i z e d , a n d h o w
fossils are b r o u g h t t o t h e surface b y e a r t h m o v e m e n t s a n d e r o s i o n . T h e se-
q u e n c e of e v e n t s is described in t h e text.
want to write about Brachiosaurus brancai (the best-known species)

would also be true of Brachiosaurus altithorax.
There are m a n y kinds of reptiles, and table 1.3 shows how they are
classified. The dinosaurs are put in the group called the Archosauria,
which also includes the crocodiles. T h e m e m b e r s of this group can be
recognized by their teeth and by the holes in their skulls. Their teeth
are set in sockets, not just stuck to the jaw like other reptile teeth.
There are more holes in the sides of their skulls than in any other
group of reptiles (figure 1.6). Holes 1 and 2 (for the nostril and eye) can
be found in the skulls of all reptiles. T h e y are the only holes in the
sides of the skulls of turtles and other Anapsida, but other reptiles have
one or both of holes 3 and 4 behind the eye. Hole 5 in front of the eye
is peculiar to archosaurs, but crocodiles do not have it. In addition
crocodiles and some other archosaurs have hole 6 in the lower jaw.
Few people would confuse crocodiles w i t h dinosaurs: crocodiles (in-
cluding alligators, etc.) are crocodile-shaped, and they have m a n y dis-
tinctive anatomical features. Pterosaurs, the other well k n o w n archo-
saurs, are also obviously different from dinosaurs: they are the winged
reptiles that are the subject of chapter 8. There were also some early
archosaurs called thecodonts, which are less obviously different from
the smaller dinosaurs, but the joints of the legs show an important
difference. Thecodont leg joints show that they m u s t have walked like
crocodiles, with their feet well out on either side of the body. Dino-
saurs walked like birds and m a m m a l s , w i t h their feet well under the
body (figure 3.5).
N o w I will review the m a i n groups of dinosaurs, introducing most
of the genera that will feature in later chapters. I will describe their
TABLE 1.1. T h e divisions of geological t i m e .
Date of Beginning
Era Period (million years ago)
Cenozoic Quaternary 1
Tertiary 65
Mesozoic Cretaceous 140

Jurassic 190
Tnassic 230
Palaeozoic (six p e r i o d s ) 570
Precambrian
N O T E : D i n o s a u r s lived during the M e s o z o i c era.

appearance, making particular m e n t i o n of their teeth and other evi-

dence of what they ate. T h e dinosaur groups and the generic n a m e s of
examples are shown in table 1.2.
The group of dinosaurs called the theropods seem, from the shapes
of their teeth, to have been flesh eaters. This sets t h e m apart from all
the other dinosaurs, which ate plants. The theropods had relatively small
fore legs and presumably walked on their hind legs. Compsognathus
(figure 1.2) was one of the smallest of t h e m . Its small, sharp teeth look
suitable for eating insects and small vertebrates, and I have already
mentioned a lizard found in one as fossil stomach contents. It lived
late in the Jurassic but there had been similar small theropods since
the Triassic. T h e ancestors of all the dinosaurs were probably rather
like Compsognathus.
Allosaurus (figure 1.7) lived at the same time but was enormously
larger. Its big teeth had serrated edges like steak knives, and look suit-
able for slicing through flesh. It probably attacked large plant-eating
dinosaurs: in a later paragraph I describe one that seems to have been
eaten by it.
Tyrannosaurus (figure 1.8) is a very late dinosaur, from the end of
the Cretaceous. It is the biggest k n o w n flesh-eating dinosaur, with 15
centimeter steak-knife teeth. It has big hind legs and small front ones
like other theropods, only more so. Its tiny front legs look useless.
TABLE 1.2. Classification of dinosaurs mentioned in this book and

the periods in which they lived.
Early
Late lurassic Cretaceous Late Cretaceous
SAURISCHIANS
thcropods Allosaurus Tyrannosaurus
Compsogna-
thus
sauropods Brachiosaurus Brachiosaurus
Diplodocus
Apatosaurus
ORNITHISCHIANS
ornithopods Iguanodon Anatosaurus
Parasaurolophus
ceratopians Psittacosaurus Triceratops
Protoceratops
pachycephalosaurs Stegoceras
stegosaurs Stegosaurus
ankylosaurs Euoplocephalus
The next major group, the sauropods, includes the largest dinosaurs.
The best k n o w n ones lived late in the Jurassic period. Diplodocus (fig-
ure 1.9) was extraordinarily long (27 meters) but was m u c h skinnier
and presumably m u c h lighter than Brachiosaurus and the other giants
that have already been mentioned. It had a very long neck and an ex-
ceedingly long tail (I discuss their functions in chapter 5), with a rel-
atively short body between. Its head looks small compared to the rest
of the animal but is about the size of a rhinoceros head. Across the
front of each jaw it has a row of tall thin teeth, like the teeth of a huge
comb, which look suitable for plucking leaves and shoots from trees
and other plants. It cannot have eaten flowering plants (which did not
evolve until the Cretaceous) and probably ate the leaves of conifers,
the commonest trees of its time. Another sauropod has been found with
fossil stomach contents which consist (rather surprisingly) of frag-
m e n t s of woody twigs of about one c e n t i m e t e r diameter. Any leaves
that were eaten with the twigs have been digested or decayed.
Large leaf-eating m a m m a l s such as giraffes pluck leaves w i t h their
tongues and front teeth, and grind t h e m w i t h their back teeth, but
Diplodocus has no back teeth and its front ones seem unsuitable for
grinding food. It has been suggested that sauropods swallowed stones
and kept t h e m in their stomachs. M o v e m e n t s of a muscular stomach
wall could have rubbed the stones together, grinding any food that was
in the stomach at the time. Plant-eating birds hold stones in their giz-
zards and use t h e m in this way: for example, ostriches have up to 900
grams (two pounds) of pebbles.
Apatosaurus (figure 1.7) was shorter but stouter than Diplodocus.
There used to be confusion about its name, because a fossil named
Apatosaurus and another originally named Brontosaurus turned out to
be identical. Apatosaurus is accepted as the correct scientific name,
but people still speak informally of brontosaurs. One fossil of Apato-
TABLE 1.3. How reptiles are classified.
class Reptilia
subclass Anapsida earliest reptiles, turtles, etc
subclass Lepidosauria lizards, snakes etc.
subclass Archosauria thecodonts
crocodiles
dinosaurs
p t e r o s a u r s (ch. 8 )
subclass Euryapsida p l e s i o s a u r s (ch. 9). e t c .
subclass Ichthyopterygia i c h t h y o s a u r s (ch. 9 )
subclass Synapsida the ancestors of m a m m a l s
F I G U R E 1.6. S k u l l s of (a) a c r o c o d i l e ; (b) a s a u r i s c h i a n d i n o s a u r [Compsog-

nathus); a n d (c) an o r n i t h i s c h i a n d i n o s a u r [lguanodon). S k u l l s (a) a n d (c) h a v e
b e e n d i s t o r t e d t o s h o w h o l e 3 , w h i c h i s i n t h e roof o f t h e s k u l l a n d w o u l d b e
h i d d e n in a true side view.
saurus has parallel scratches on its ribs, spaced about the same dis-
tance apart as the teeth of Allosaurus. Beside it was found a broken-
off Allosaurus tooth. It seems that an Allosaurus fed on the carcass,
but there is nothing to show w h e t h e r it killed the sauropod (which was
m u c h larger than it could have been) or found the sauropod already
dead.
Sauropods had elephant-like fore feet that would not have been much
use for handling things. It seems obvious that they walked on all fours
(unlike theropods), and there are fossil footprints that seem to confirm
this. (They are described in chapter 3). However, the shortness of the
fore legs of Diplodocus and Apatosaurus suggests that they may have
evolved from bipedal ancestors. Also, there are early members of the
FIGURE 1.7. Apatosaurus, Allosaurus (the biped), and a large African ele-
phant.
INTRODUCING THE DINOSAURS I 1
F I G U R E 1.8. Tyrannosaurus, f r o m N e w m a n 1 9 7 0 , a n d a b a s k e t b a l l n e t . T h e
m o u t h o f t h e n e t i s 3 . 0 5 m e t e r s (10 ft] f r o m t h e f l o o r .
group that may have been bipeds. Brachiosaurus (figure 1.1) was a dif-
ferently proportioned sauropod, with very long front legs and shorter
hind ones. Its vertebrae fit together best w i t h the neck nearly vertical,
suggesting that that was how the neck was carried. T h e long front legs
and near-vertical neck would have enabled it to reach high branches,
to feed like a giraffe. Its teeth were broader than Diplodocus' teeth but
still seem suitable only for plucking off leaves.
The theropods and sauropods together form the Saurischia, one of
the two main groups of dinosaurs. The remaining dinosaurs (all of t h e m
plant eaters) form the Ornithischia. T h e most obvious differences be-
tween saurischians and ornithischians are in their jaws and hips. Com-
pare the skull of Iguanodon (an ornithischian, figure 1.6c) w i t h that of
Compsognathus (a saurischian, figure 1.6b). T h e ornithischians had no
front teeth and presumably had horny beaks, like the beaks of birds
and turtles, on the fronts of their jaws. (I have already mentioned the
parrot-like beak of Psittacosaurus, another ornithischian.) Another pe-
F I G U R E 1.9. Diplodocus a n d a S a n F r a n c i s c o c a b l e c a r . T h e d i n o s a u r , w h i c h
is 25 m e t e r s long, is d r a w n from a m o d e l sold by the N a t u r a l H i s t o r y M u s e u m ,
London.
culiarity of ornithischians is an extra bone (the predentary) at the front

of the lower jaw.
Figure 1.10 shows the pelvic girdles of Tyrannosaurus (a saurischian)
and Psittacosaurus (an ornithischian). You can see how these bones fit
into the skeleton, in the hip region, by referring back to figures 1.8 and
1.4. In saurischians the pubis points forward and the ischium back-
ward, but in ornithischians the pubis also has a backward extension.
The first of the ornithischians that I will describe belong to the group
called the ornithopods. Anatosaurus (figure 1.11) and Iguanodon (figure
3.6) are examples. They m u s t have carried at least most of their weight
on their big hind legs, but m a y s o m e t i m e s have rested their front feet
on the ground as well. Anatosaurus has a broad duck-like beak but
Iguanodon has a deeper, narrower one. Both have impressive batteries
of grinding teeth (a striking difference from sauropods) and m u s t have
chewed their food like cattle. It has been suggested that they may have
had cheeks enclosing the sides of their m o u t h s to prevent half-chewed
food from falling out. M a m m a l s have cheeks, but modern reptiles do
not.
Teeth that were simply pressed together would crush food, but to
grind food they m u s t slide over each other. Horses and cattle grind by
moving their lower jaws from side to side, but ornithopod jaws worked
differently. T h e lower jaws moved straight up and down but the upper
jaws were hinged along their upper edges and splayed apart when the
teeth pressed together (figure 1.12). T h u s the teeth slid over each other,
grinding the food rather than merely crushing it. No one has seen this
m e c h a n i s m working (there is nothing like it in living animals), but
FIGURE 1.10. P e l v i c g i r d l e s o f a s a u r i s c h i a n {Tyrannosaurus) a n d a n o r n i t h i s -

chian {Psittacosaurus).
F I G U R E 1.11. Anatosaurus, f r o m G a l t o n 1 9 7 0 , c h a s i n g a P o r s c h e . T h e d i n o -
s a u r i s 8.9 m e t e r s l o n g a n d t h e c a r 4 . 3 m e t e r s .
study of the structure of the jaws and their joints suggests that they
must have worked in this way. Further evidence has been obtained by
looking at the worn surfaces of the teeth through a microscope: the
fine scratches on t h e m run across the teeth, as they should if the jaws
moved as suggested.
The ceratopians or horned dinosaurs were another important group.
Psittacosaurus (figure 1.4) is a ceratopian, but it is unusual in two ways:
it had no horns, and it seems to have walked on two legs whereas other
ceratopians walked on all four. Triceratops (figure 1.13) is more typical,
and is one of the largest ceratopians. It has two long horns, one over
each eye, and a short one on its snout. It also has a great frill of bone
extending back from the skull over the neck. Some other ceratopians
have even longer frills. I discuss these horns and frills in chapter 6.
Ceratopians had impressive batteries of back teeth, arranged differ-
Iguanodon Triceratops
F I G U R E 1.12. D i a g r a m m a t i c s e c t i o n s t h r o u g h t h e j a w s o f a n o r n i t h o p o d a n d
a ceratopian. E n a m e l coatings on tooth surfaces arc represented by thick lines.
Arrows show how the jaws moved.
FIGURE 1.13. Triceratops ( c e n t e r ) a n d Stegosaurus (right), d r a w n f r o m m o d e l s

sold by the N a t u r a l H i s t o r y M u s e u m , London. T h e plant is a cycad, such as
Triceratops m a y h a v e e a t e n . A l a r g e B l a c k r h i n o c e r o s i s a l s o s h o w n : i t i s 3 . 5
m e t e r s long, i n c l u d i n g t h e tail.
ently from the teeth of ornithopods. T h e upper and lower teeth did not
meet crown-to-crown w h e n the m o u t h closed, so they could not crush
or grind food. Instead, the lower teeth came up close inside the upper
ones (figure 1.12). We can be sure that they worked like this because
the worn surfaces of the teeth are vertical. T h u s the rows of teeth moved
like the blades of shears. They would have been very effective for chop-
ping up fibrous plant food, perhaps leaves of palms or cycads.
T h e pachycephalosaurs are a rather rare group of bipedal ornithis-
chains. T h e y have astonishingly thick skull roofs: some are 23 centi-
meters thick of solid bone. I have written about the function of this
thickening in chapter 6.
T h e stegosaurids, another group of ornithischian dinosaurs, have a
row of big bony plates along their backs (figure 1.13). I m e n t i o n a pos-
sible function of these plates in chapter 7. Stegosaurids have long hind
legs and short fore legs, but seem to have walked on all fours. Their
heads are relatively small and their teeth are m u c h less impressive than
those of ornithopods and ceratopians. T h e y have formidable spines on
their tails.
T h e ankylosaurs are the last group of dinosaurs in this list. They had
short legs and broad bodies, and walked on all fours. They had thick
plates of bone embedded as armour plating in their skin, as if they were
gigantic armadilloes. Some had club-like l u m p s of bone at the ends of
their tails. I discuss t h e m in chapter 5.
Dinosaurs lived on most of the earth's land mass throughout the
Mesozoic era, but different dinosaurs lived in different places, at dif-
ferent times. Tyrannosaurus never tried its strength against Apatosau-
rus, which had been extinct for 70 million years before Tyrannosaurus
evolved. Compsognathus, which lived in Europe, probably never met
Allosaurus, which lived in the western United States, although they
overlapped in t i m e . O t h e r groups of famous dinosaurs m u s t have met.
Allosaurus, Apatosaurus and Stegosaurus were all living in North
America at the same time, late in the Jurassic period. Seventy million
INTRODUCING THE DINOSAURS 1 5
years later, late in the Cretaceous, Tyrannosaurus, Anatosaurus, and

Triceratops were all living in N o r t h America.
Principal Sources
C h a r i g (1979) i s a n e x c e l l e n t s h o r t i n t r o d u c t i o n t o t h e d i n o s a u r s . N o r m a n (1985) i s
a r e m a r k a b l e m i n e o f i n f o r m a t i o n , b u t u n f o r t u n a t e l y i n c l u d e s n o references t o t h e
m o r e s p e c i a l i z e d scientific l i t e r a t u r e . B a k k e r (1986) is an i d i o s y n c h r a t i c a c c o u n t of
t h e d i n o s a u r s by a s c i e n t i s t w h o s e w o r k h a s s t i m u l a t e d a g r e a t deal of i n t e r e s t a n d
c o n t r o v e r s y . C z e r k a s a n d O l s o n (1987) s h o w s h o w s c i e n t i s t s ' v i e w s a b o u t d i n o s a u r s
h a v e c h a n g e d i n r e c e n t years. N o r m a n a n d W e i s h a m p e l (1985) d e s c r i b e t h e j a w
m e c h a n i s m of o r n i t h o p o d s . Paul (1988) c o m p a r e s t h e sizes of t h e largest d i n o s a u r s .
T h e o t h e r references i n t h e list t h a t f o l l o w s a r e s o u r c e s for i l l u s t r a t i o n s .
Bakker, R. T. 1986. The Dinosaur Heresies. N e w Y o r k : M o r r o w .

C h a r i g , A. 1979. A New Look at the Dinosaurs. L o n d o n : British M u s e u m ( N a t u r a l
History).
C o o m b s , W . P . 1980. J u v e n i l e c e r a t o p s i a n s f r o m M o n g o l i a : T h e s m a l l e s t k n o w n di-
n o s a u r s p e c i m e n s . Nature 2 8 3 : 3 8 0 - 3 8 1 .
C z e r k a s , S. J. a n d E. C. O l s o n . 1987. Dinosaurs Past and Present. S e a t t l e : U n i v e r s i t y
of W a s h i n g t o n Press.
G a l t o n , P. M. 1970. T h e p o s t u r e of h a d r o s a u r i a n d i n o s a u r s , journal of Palaeontology
44:464-473.
N e w m a n , B. H. 1970. S t a n c e a n d gait in t h e f l e s h - e a t i n g d i n o s a u r Tyrannosaurus.
Biological lournal of the Linnean Society 2 : 1 1 9 - 1 2 3 .
N o r m a n , D. B. 1985. The Illustrated Encyclopedia of Dinosaurs. L o n d o n : S a l a m a n -
der Books.
N o r m a n , D . B . a n d D . B . W e i s h a m p e l . 1985. O r n i t h o p o d feeding m e c h a n i s m s : T h e i r
b e a r i n g on t h e e v o l u t i o n of h e r b i v o r y . American Naturalist 1 2 6 : 1 5 1 - 1 6 4 .
O s b o r n , H. F. 1924. Psittacosaurus a n d Protiguanodon: T w o L o w e r C r e t a c e o u s ig-
u a n o d o n t s from M o n g o l i a . American Museum Novitates 1 2 7 : 1 - 1 6 .
Paul, G. S. 1988. T h e b r a c h i o s a u r g i a n t s of t h e M o r r i s o n a n d T e n d a g u r u . Hunteria
2(3): 1 - 1 3 .
II
Weighing Dinosaurs
of dinosaurs in chapter 1 were based on fossil bones,

r-i-i H E D R A W I N G S
.L fitted together to form complete skeletons. These skeletons tell

us how large dinosaurs were. There is no reason to suspect that the
sizes of bones have been changed by the processes of fossilization. There
m a y of course be doubts about the size of the animal if bones are miss-
ing. For example, there are doubts about how m a n y bones are missing
from the tail of Tyrannosaurus, which one palaeontologist made 3.7
meters (12 ft) longer than another. There are also dangers of error if a
complete skeleton has been assembled from bones of different individ-
uals: animals of different sizes may be combined, producing an ill-pro-
portioned skeleton. However, m a n y of the best-known skeletons have
been made from bones found together, apparently from one individual
animal.
M e a s u r e m e n t s of skeletons tell us how long and how tall dinosaurs
were, but do not directly tell us what they weighed. We might want
to know, for at least two reasons. First, the weight (or mass) of an an-
imal seems a fairer summary of its size than any measurement of height
or length. Giraffes are taller than elephants and some pythons are longer
but it is the elephant, the heaviest of the three, that impresses us as
the largest. Second, some dinosaurs were so large as to make us wonder
how easily their legs could have supported their weight. Were they as
active as modern reptiles or were they lumbering monsters, barely able
to move about? Were they perhaps so heavy that they had to live partly
submerged in water, which would help to support t h e m by buoyancy?
To tackle questions like these, we want to know just how heavy the
big dinosaurs were.
There is an awkward difference between the scientific meaning of
the word " w e i g h t " and its use in everyday language. Non-scientists
often give the " w e i g h t s " of objects in pounds or kilograms, but sci-
WEIGHING DINOSAURS I 7
entists measure the quantity that they call weight in newtons. To us,
weight means the force exerted on an object by gravity. It is measured
in newtons, the unit of force. " M a s s " is our n a m e for the quantity that
we measure in kilograms. (We don't use pounds.) Weight is calculated
by multiplying mass by the gravitational acceleration g which is about
10 m e t e r s / s e c o n d : that means that the speed of a falling body, un-
2
hindered by air resistance or anything else, increases by 10 meters per

second every second. If a body has a mass of m kilograms, its weight
on earth is 10m newtons.
Elephants, rhinoceroses, and (we will see) the larger dinosaurs have
masses of several thousand kilograms. This makes it convenient to give
their masses in tonnes (metric tons). T h e tonne is 1,000 kilograms or
2,205 pounds, almost the same as the commercial (long) ton of 2,240
pounds.
Modern animals can be weighed, but the masses of dinosaurs can
only be estimated from other m e a s u r e m e n t s . It seems sensible to give
the masses of some large modern animals, as bases for comparison,
before going on to the extinct ones.
Though it is possible to weigh modern animals, the biggest ones
present problems. Domestic and zoo animals can be driven onto weigh-
bridges like the ones used for weighing vehicles, but large wild animals
are difficult to transport (alive or dead) to such facilities. Most available
masses of large wild animals are of specimens shot and weighed in the
field. Some were shot in the course of population control or scientific
research, and others by hunters. A team culling a hippopotamus pop-
ulation used a two-tonne spring balance m o u n t e d on a hydraulic hoist
on the back of a truck, but large animals usually have to be cut up and
weighed in pieces. Some blood and other fluids are lost in the process,
but the loss is generally no more than 3 percent of body mass.
Some masses of modern animals are given in table 2.1. All (except
the humans) were wild animals killed in the field. Elephants continue
growing throughout life and the masses given for t h e m are the limits
that they seem to approach in old age. T h e other masses are m e a n s
from groups of adults. The Blue whale is the largest of the whales. T h e
African elephant grows larger than the Indian elephant, which is not
included in the table. The Black rhinoceros, however, is not the largest
of the rhinoceroses. I have unfortunately not managed to find reliable
masses for the White rhinoceros, which is said to surpass three tonnes.
The lion is a large terrestrial carnivore, but not the largest. Tigers grow
a little heavier, and the larger species of bear exceed half a tonne.
Now we return to the dinosaurs. H o w can we weigh them? In most
cases, we have only their bones to guide us. We cannot estimate the
masses of the living animals from the masses of their bones, because
1 8 WEIGHING DINOSAURS
these have been altered by the processes of fossilization. They have lost
water and protein and become impregnated w i t h minerals.
We want to estimate not just the mass of the skeleton but its mass
with all the guts, flesh, and skin that went w i t h it. It seems easiest to
base our calculations on models of the animals as we think they would
have looked in life. Many such models have been made. The best com-
mercially available ones that I k n o w are plastic models at 1:40 scale,
sold by the Natural History Museum, London. These seem to have been
made carefully and accurately. I have checked m a n y of their dimen-
sions and find that they are indeed about 1:40 of the corresponding
dimensions of the best-known fossil skeletons of the species they rep-
resent. I have used these models for estimating dinosaur masses. How-
ever, I once wanted to estimate the masses of moas (extinct ostrich-
like birds from N e w Zealand) and could not find suitable models. I
modeled the m a i n features of the skeleton to scale in soldered wire,
and used modeling clay to build up the flesh around it.
T h e masses of models depend on the densities of the materials used
to m a k e them, which may not be the same as the tissues of the ani-
mals. For this reason, the first stage in finding dinosaur masses is to
measure the volume, not the mass, of the model.
Edwin Colbert, a distinguished U.S. paleontologist, measured the
volumes of models in this way. He put a model in a box and packed
sand around it until the box was filled to the brim. T h e n he removed
the model, being careful to spill none of the sand, and added more sand
until the box was full again. This extra sand had the same volume as
the model.
TABLE 2.1. Masses (in tonnes) of some modern animals.
Males Females
Blue whale, Balaenoptera musculus 91 110
African elephant, Loxodonta africana 5.45 2.77
Hippopotamus, Hippopotamus amphibius 2.52 2.13
Black rhinoceros, Diceros bicornis 1.17 1.08
Eland, Taurotragus oryx 0.84 —
African buffalo, Syncerus caffer 0.75 —
L i o n , Panther a leo 0.18 0.15
Human, Homo sapiens 0.07 0.05
S O U R C E S : w h a l e : N i s h i w a k i 1950; e l e p h a n t : Laws 1966; others: M e i n e r t z h a g e n 1938.

WEIGHING DINOSAURS 19
Colbert's method was difficult to perform accurately because it de-

pended on the sand being leveled off precisely at the top of the box.
He used it in preference to the m e t h o d that I used later, because he
did not want to get his valuable plaster models wet. For m e a s u r e m e n t s
on plastic models, I used a m e t h o d that depends on Archimedes' Prin-
ciple, the principle of buoyancy. When an object is submerged in water,
the water exerts an upward force on it. This force (called the upthrust)
equals the weight of as m u c h water as would have the same v o l u m e
as the submerged body.
The diagram (figure 2.1) shows the m e t h o d . T h e model is suspended
by a thread from one arm of a beam balance, w i t h a metal weight hang-
ing from its tail. (The weight is unnecessary if the model is denser than
water.) It hangs in a tall jar, not touching the b o t t o m or the sides. Ini-
tially, the weight (if any) is submerged in water but the whole model
is above the water. Enough weights are put on the pan to balance the
system. Then water is added until the model is completely submerged.
The upthrust of the water on the model puts the system out of balance
and weights m u s t be removed from the pan, to restore the balance. For
example, when the experiment was done w i t h a model of Brachiosau-
rus, weights totaling 728 grams had to be removed. This showed that
the volume of the model equaled the volume of 728 grams water: it
was 728 cubic centimeters.
The volume of the model m u s t be scaled up to get the v o l u m e of
the dinosaur. This particular model was 1:40 scale. Therefore the di-
nosaur was 40 times as long, 40 times as wide and 40 t i m e s as high
as the model. Its volume was 40 x 40 x 40 = 64,000 t i m e s the volume
of the model: it was 64,000 x 728 = 46,600,000 cubic centimeters or
46.6 cubic meters.
To get the mass of the dinosaur from its volume, we m u s t estimate
its density. Most animals have about the same density as water. They
either just float in water, with very little projecting above the surface,
or just sink. Among living animals, crocodiles probably give the best
indication of the probable density of dinosaurs. N o t only are they among
the largest modern reptiles, but they are believed to be quite closely
related to the dinosaurs. Hugh Cott, a British zoologist, measured the
densities of nine dead Nile crocodiles and got a m e a n value of 1,080
kilograms per cubic meter. This is quite a lot greater than the density
of water, which is 1,000 kilograms per cubic meter. However, the lungs
of the dead crocodiles were probably deflated, and live ones would have
been less dense. Cott observed that crocodiles often float in water w i t h
only the nostrils and top of the head above the surface. They m u s t then
be very slightly less dense than water. He also observed that they
sometimes rest motionless on the b o t t o m s of rivers, w h e n they m u s t
20 WEIGHING DINOSAURS
FIGURE 2.1. A p p a r a t u s for m e a s u r i n g t h e v o l u m e s o f m o d e l d i n o s a u r s .
be denser than water. Plainly, crocodiles can vary their density by in-
flating and deflating their lungs, but they are probably always very close
to the density of water. I will therefore assume that dinosaurs had a
density of 1,000 kilograms per cubic meter. Colbert assumed a smaller
density, based on m e a s u r e m e n t s on small reptiles that were probably
inaccurate.
T h e volume of Brachiosaurus was estimated to be 46.6 cubic meters.
If its density was 1,000 kilograms per cubic meter its mass was 46,600
kilograms, or 46.6 tonnes. This is colossal. It is about nine times the
mass of a large male African elephant, the largest modern terrestrial
animal. It is only half the mass of a large Blue whale, but whales live
submerged in water which supports their weight by buoyancy.
T h e m e t h o d of calculating masses from the volumes of models re-
quires a fairly complete skeleton, unless the model maker is prepared
to risk guessing the sizes of missing parts. An alternative approach us-
ing the dimensions of just a few selected bones was used in a recent
international project. T h e collaborators were J. F. Anderson, a U.S. zo-
ologist with a long-standing interest in the sizes of bones of different-
sized modern animals; A. Hall-Martin, w h o works at Kruger National
Park, South Africa; and Dale Russell, a Canadian dinosaur specialist.
Chapter 3 will show that dinosaurs stood and moved m u c h more
WEIGHING DINOSAURS 2 1
like m a m m a l s than like crocodiles, lizards, and tortoises. T h e Ander-

son team chose to study m a m m a l bones to look for rules that would
enable t h e m to estimate dinosaur masses. T h e University of Florida,
where Anderson works, has a large collection of m a m m a l skeletons
with records of the masses of the intact animals. T h a t collection in-
cludes few really large m a m m a l s , but Hall-Martin was able to measure
the bones of animals shot for other purposes in Kruger National Park.
They chose to study major leg bones, which are often well preserved
in otherwise incomplete fossils. T h e y could have used the lengths of
the bones, but this might have led to errors due to some animals hav-
ing spindly legs and others s t u m p y ones. An extreme case will show
how serious this could be. Figure 2.2 shows two birds measured in Kenya
by Professor Geoffrey Maloiy and m e . T h e Secretary bird was slightly
lighter than the Ground hornbill, but its leg bones were up to twice as
long. The diameters and circumferences of leg bones seem to be better
indicators of body mass, than are their lengths.
The Anderson team chose to study the circumferences of the hu-
merus and femur, the bones of the upper parts of the legs (figure 2.3).
They measured the circumferences of these bones where they were least,
about half way along the shafts of the bones, and (in their studies of
quadrupeds) added the t w o circumferences together. T h e y could have
used the circumference of either bone alone, but that would have led
F I G U R E 2.2. (a) a 3 . 9 - k i l o g r a m S e c r e t a r y b i r d [Sagittarius serpentarius a n d (b)

a 4 . 2 - k i l o g r a m G r o u n d h o r n b i l l [Bucorvus leadbeateri). T h e i r m a s s e s a r e a b o u t
t h e s a m e but t h e l e n g t h s of their leg b o n e s are very different.
22 WEIGHING DINOSAURS
FIGURE 2.3. The skeleton o f Brachiosaurus brancai, showing the n a m e s of

some bones.
to errors due to different quadrupeds supporting different fractions of

their weight on their fore and hind legs. Apatosaurus seems to have
carried most of its weight on its hind legs, and has a thick femur and
a t h i n n e r h u m e r u s . Brachiosaurus seems to have carried a bigger frac-
tion of its weight on its fore legs, and the two bones are more nearly
equal in circumference. Any formula based on one bone alone would
probably overestimate the m a s s of one of these dinosaurs, and under-
estimate the other.
Figure 2.4 shows the total of h u m e r u s and femur circumferences
plotted against body mass, for quadrupedal m a m m a l s . The scale of the
graph has been chosen so that an exceptionally large elephant can just
be fitted in, but that leaves most of the points for other animals squashed
into the b o t t o m left corner. It has been impossible to show the data
clearly for anything smaller t h a n a 29-kg baboon.
Figure 2.5 shows the same data (and more) plotted in a different way.
T h e b o t t o m scale has been distorted so that the distance from the 10-
gram mark to the 100-gram m a r k is the same as from 100 grams to
1,000 grams (1 kilogram) or even from 10 tonnes to 100 tonnes. The
scale up the side has been distorted in the same way. More precisely,
WEIGHING DINOSAURS 23
distances along both scales are proportional to the logarithms of the

numbers they represent, not to the numbers themselves. This has made
it possible to show data for the whole range of sizes from mice to di-
nosaurs.
The solid black points in figure 2.5 are data for quadrupedal m a m -
mals. They form a more-or-less straight band. T h e line has been fitted
to t h e m by statistical analysis. It is the best straight line that can be
got from this set of data, for estimating body mass from bone circum-
ferences. Its equation (written in the m o s t convenient form for the pur-
pose] is
body mass in kg = 0.000084 (total of circumferences in m m ) 2 73
(If you want to use this equation to estimate the body mass of a par-
ticular animal from its h u m e r u s and femur circumferences, you will
have to use the y" button on your calculator to raise the total circum-
ference to the 2.73 power.)
All the black points lie close to the line, which suggests that it should
be possible to estimate the masses of m a m m a l s rather accurately, from
Body m a s s (tonnes)
F I G U R E 2.4. A g r a p h s h o w i n g ( h u m e r u s c i r c u m f e r e n c e p l u s f e m u r c i r c u m f e r -
e n c e ) p l o t t e d a g a i n s t b o d y m a s s , for v a r i o u s q u a d r u p e d a l m a m m a l s . D a t a f r o m
A n d e r s o n e t al. 1 9 8 5 .
F I G U R E 2.5. T h e s a m e d a t a a s i n figure 2.4 re-plotted o n l o g a r i t h m i c scales,
w i t h a d d i t i o n a l p o i n t s , i n c l u d i n g o n e for Brachiosaurus.
the circumferences of their bones. However, appearances can be de-

ceptive. Imagine that we do not know the masses of the two largest
m a m m a l s in the graph, and want to estimate t h e m from their bone
circumferences. T h e estimates that the equation would give us are 1.2
tonnes for the 2.0-tonne hippopotamus, and 9.0 tonnes for the 5.9-tonne
elephant. We m u s t expect errors as bad as this, or worse, if we use the
equation for estimating dinosaur masses.
T h e circumferences of the h u m e r u s and femur of Brachiosaurus are
654 and 730 millimeters, giving a total 1384 millimeters. The mass of
the same individual Brachiosaurus has already been estimated, from
the volume of a model, to be 47 tonnes. These data are represented by
the hollow circle in figure 2.5, which lies well below the m a m m a l line.
The line predicts a mass of only 32 tonnes, for the observed circum-
ferences, but the discrepancy is no worse t h a n the examples of the hip-
popotamus and elephant led us to expect.
T h e line is based on quadrupedal m a m m a l s , and it seems reasonable
to apply it to quadrupedal dinosaurs. It would be a mistake to expect
it to give accurate estimates of body mass, but even rough estimates
are interesting. For bipeds, a different line is needed. The Anderson
team produced a biped equation by modifying the quadruped one. They
used only the femur circumference (because bipeds walk only on their
hind legs) and adjusted the factor in the equation to m a k e the predicted
mass for one particular dinosaur m a t c h the mass that had been esti-
mated from a model. Their biped equation is
body mass in kg = 0.00016 (femur circumference in mm) ™. 2

WEIGHING DINOSAURS 2S
This underestimates the masses of kangaroos and overestimates the

masses of ostriches.
Table 2.2 shows the masses of dinosaurs estimated from the vol-
umes of models by Colbert and myself, and calculated from their equa-
tions by the Anderson group. For each of the species in the table, at
least two estimates of mass have been made. For some of them, the
estimates agree well, but for others there are big discrepancies. In the
worst cases {Diplodocus and Brachiosaurus) the largest estimate is about
three times the smallest.
Some of the discrepancies can be explained. Photographs in Colbert's
paper show that his model of Stegosaurus was skinnier than the one
I used, and his model of Triceratops was more portly than mine. In
each case, one model may be more realistic than the other, but it is
hard to say which. Even if a model is based on accurate skeleton mea-
surements, its volume depends a lot on the judgement of the modeller.
Some other discrepancies may be due to estimates being based on dif-
ferent-sized specimens of the same species.
TABLE 2.2. Masses (in tonnes) of dinosaurs.
Colbert' 1
Alexander' Anderson et al.h
(1962) (1985) (1985)
theropods
Allosaurus fragilis 2.3 1.4
Tyrannosaurus rex 7.7 7.4 4.5
sauropods
Diplodocus carnegiei 11.7 18.5 5.8
Apatosaurus louisae 33.5 — 37.5
Brachiosaurus brancai 87.0 46.6 31.6
ornithopods
Iguanodon hernissartensis 5.0 5.4 —
'Anatosaurus' copei 3.4 — 4.0
stegosaurs
Stegosaurus ungulatus 2.0 3.1 —
ccratopians
Styracosaurus alhertensis 4.1 4.1
Triceratops 'prorsus' 9.4 —
N O T E : T h e Colbert e s t i m a t e s , a s originally published, w e r e based o n a n a s s u m e d d e n s i t y o f 9 0 0

kilograms per cubic meter. T h e y h a v e b e e n adjusted to a d e n s i t y of 1,000 k i l o g r a m s per cubic
meter, the same as for the Alexander values. T h e Anderson values for quadrupeds are also slightly
larger than t h o s e g i v e n in the original paper, b e c a u s e an error in the e q u a t i o n has been corrected.
T h e Brachiosaurus e s t i m a t e in the Alexander c o l u m n d o e s not appear in A l e x a n d e r 11985] but
w a s obtained by the s a m e m e t h o d .
' E s t i m a t e d from the v o l u m e s of m o d e l s .
b
E s t i m a t e d from the c i r c u m f e r e n c e of bones.
2 6 WEIGHING DINOSAURS
It is hard to decide w h i c h of the mass estimates to prefer, in table

2.2. You might prefer the estimates based on models because they take
account of the whole skeleton, not just the leg bones. Alternatively
you might prefer the Anderson estimates because they do not depend
on the judgment of a modeler. I am inclined to prefer the estimates
from models.
Table 2.2. shows that there is uncertainty about the masses of di-
nosaurs. However, it is quite clear that the largest dinosaurs were ex-
ceedingly heavy. Compare this table w i t h table 2.1, which gives the
masses of modern animals. Whichever estimate you prefer, the bulky
sauropods [Apatosaurus and Brachiosaurus) were many times as heavy
as the biggest African elephants. T h e very long but slender Diplodocus
was probably also m u c h heavier than elephants: the sketch of one be-
side a cable car (figure 1.9) m a k e s it hard to believe the very low An-
derson estimate of its mass. Triceratops, the big ceratopian, seems to
have been at least as heavy as most elephants. T h e biggest bipeds, both
carnivores such as Tyrannosaurus and herbivores such as Iguanodon,
were in the same range of masses, and m a n y other dinosaurs were at
least as heavy as an adult hippopotamus.
Finally, please r e m e m b e r that m a n y dinosaurs were a great deal
smaller than the ones in the table. T h e young Psittacosaurus in figure
1.2 is smaller than the pigeon.
Principal Sources
T h e e s t i m a t e s o f d i n o s a u r m a s s e s c o m e from t h e p a p e r s o f C o l b e r t , 1962, A l e x a n d e r
1985, a n d A n d e r s o n e t al. 1985. T h e o t h e r p a p e r s i n t h e list b e l o w are s o u r c e s for
t h e m a s s e s of m o d e r n a n i m a l s g i v e n in t a b l e 2.1 a n d for figure 2 . 3 .
A l e x a n d e r , R. M c N . 1985. M e c h a n i c s of p o s t u r e a n d gait of s o m e large d i n o s a u r s .

Zoological lournal of the Linnean Society 8 3 : 1 - 2 5 .
A n d e r s o n , J. F., A. H a l l - M a r t i n , a n d D. A. R u s s e l l . 1985. Long b o n e c i r c u m f e r e n c e
a n d w e i g h t in m a m m a l s , birds a n d d i n o s a u r s , lournal of Zoology (A) 2 0 7 : 5 3 - 6 1 .
Colbert, E. H. 1962. T h e w e i g h t s of d i n o s a u r s . American Museum Novitates 2 0 7 6 : 1 -
16.
J a n e n s c h , W. 1950. D i e S k e l c t t r e k o n s t r u k t i o n v o n Brachiosaurus brancai. Palaeon-
tographica s u p p l e m e n t 7 : 9 5 - 1 0 3 .
L a w s , R. M. 1966. Age c r i t e r i a for t h e African e l e p h a n t , Loxodonta africana. East
African Wildlife journal 4 : 1 - 3 7 .
M e i n e r t z h a g e n , R. 1938. S o m e w e i g h t s a n d m e a s u r e m e n t s of large m a m m a l s . Pro-
ceedings of the Zoological Society of London 1938A:433-439.
N i s h i w a k i , M. 1950. On t h e b o d y w e i g h t s of w h a l e s . Scientific Reports on Whales
of the Research Institute Tokyo 4 : 1 8 4 - 2 0 9 .
Ill
Dinosaur Footprints
O F A L L that remains of dinosaurs, their footprints bring the animals

most vividly to life. Fossil bones may remind us of a rotting car-
cass, but footprints are evidence of a living, moving animal. It may
seem incredible that dinosaur footprints should be preserved. T h e
chances of an individual footprint surviving are remote, but millions
of dinosaurs made footprints every day, for millions of years. For some,
the improbable happened: they survived and were found by inquisitive
people.
Figure 3.1 shows part of a remarkable group of over 3,000 footprints,
found in a patch of rock in Australia. It seems obvious that they are
footprints but they m u s t be about 100 million years old. Geological
evidence shows that the rock was formed in the middle part of the
Cretaceous period, w h e n dinosaurs were plentiful. T h e tracks of m a n y
individuals can be followed across the rock but none show any indi-
cation of distinct fore and hind feet, so the animals m u s t have been
bipeds. The biggest prints, 64 centimeters (25 inches) long, m u s t have
been made by a dinosaur: there were no other big bipeds around. T h e
others, some of t h e m as small as chicken footprints, were probably
made by smaller dinosaurs. They are not quite the same shape as mod-
ern bird footprints, and bird fossils are rare in Cretaceous rocks.
The structure of the rock seems to show h o w the footprints got pre-
served. There are alternating layers of claystone (formed from mud) and
sandstone (formed from sand). They seem to have been laid down where
a river flowed into a lake, just as sand and m u d carried down modern
rivers get deposited as sand or m u d banks at the m o u t h . In dry periods
the lake would have shrunk leaving the m u d bank exposed to the air.
At times of flood the water would rise over the m u d again and sedi-
m e n t would settle out of it. Coarse grains would settle first, forming
sand, and fine grains would settle later, forming mud.
2K DINOSAUR FOOTPRINTS
F I G U R E 3 . 1 . D i n o s a u r footprints found at W i n t o n , Q u e e n s l a n d , from T h u l -

born and W a d e 1984.
A m u d surface was exposed in a dry period when dinosaurs walked

over it, forming footprints (figure 3.2a). T h e floods came and deposited
a layer of sand, which filled the footprints (figure 3.2b). More layers of
sediment were deposited, t i m e passed and eventually the mud and sand
became rock. The footprints had been impressed in m u d and filled with
sand, but now they were impressions in claystone, filled with sand-
stone. Fortunately the t w o kinds of rock do not stick together very
firmly. Slabs of sandstone can be levered off leaving the claystone un-
damaged (figure 3.2c). After some of the prints had been found by a
happy chance it was not too difficult to expose the rest. Some sand-
stone got left behind in the hollows of the prints, but it was possible
to clean it out w i t h an awl.
We have identified the tracks as those of bipedal dinosaurs, but we
can be more specific. Both the flesh-eating theropods and the plant-
eating ornithopods were bipedal, and both had three m a i n toes. Ther-
opods had sharp claws which were probably useful for attacking prey
(figure 3.3a). Ornithopods had blunter, more hoof-like tips to their toes
(figure 3.3b). T h e biggest footprints in figure 3.1 show claw marks, so
were probably made by a theropod. Their size suggests an animal a
little smaller than Tyrannosaurus, probably about five tonnes. Some
of the smaller prints are thought to have been made by ornithopods
and some by tiny theropods.
Figure 3.4 shows some footprints found in Texas. A few of t h e m
were made by three-toed bipeds, probably theropods, which were a lit-
tle smaller than the big theropod at Winton. Most of the footprints,
however, were made by quadrupeds. There are a lot of tracks on top
of each other, giving a muddled impression, but one trackway (picked
DINOSAUR FOOTPRINTS 2 9
out in stipple) is very clear. It has huge hind footprints and smaller
fore ones. Each is the right shape to have been made by a sauropod
(figure 3.3c,d) and their size suggests an animal of 2 0 - 3 0 tonnes. T h e
biggest of the hind footprints are 76 centimeters (30 inches) long. There
are also footprints of smaller sauropods.
Even bigger footprints have been found elsewhere in Texas, with hind
prints 92 centimeters (36 inches) long.
T h e footprints in figures 3.1 and 3.4 are at exceptional sites, where
large numbers of dinosaur footprints have been found together. Less
impressive finds are quite c o m m o n . Large n u m b e r s of dinosaur foot-
prints have been found, all over the world. They can tell us a lot about
the lives of dinosaurs.
First, how did dinosaurs stand and move? Modern reptiles walk w i t h
their feet well out on either side of the body, so the lines of footprints
made by their left and right feet are well apart (figure 3.5a). Birds and
m a m m a l s walk with their feet under the body, and the lines of left and
right prints are close together (figure 3.5b,c). Dinosaur footprints show
that they walked like birds and m a m m a l s , w i t h their feet well under
the body. The shapes of dinosaur leg bones show the same thing. T h e y
would have dislocated their joints if they had tried to walk like modern
reptiles.
The footprints in figure 3.6 show an exception to the general rule.
There are large, ill-formed hind footprints (stippled) w i t h smaller fore
prints (black) on either side. T h e prints are too poor to show the shapes
of the feet that made them, but they were probably made by Iguano-
don, which was around at the t i m e . T h e hind feet were apparently un-
der the body, mammal-fashion, but the fore feet reached out to either
side. Iguanodon looks like a biped, but these tracks suggest that its
fore feet took a little of its weight.
One striking thing about dinosaur footprints is that there is seldom
any sign of marks made by the tail. Old pictures of dinosaurs show the
F I G U R E 3.2. D i a g r a m s s h o w i n g h o w t h e W i n t o n f o o t p r i n t s w e r e p r e s e r v e d .
Each d i a g r a m is a v e r t i c a l s e c t i o n t h r o u g h t h e l a y e r s of m u d (light stipple) a n d
sand (dense stipple).
30 DINOSAUR FOOTPRINTS
F I G U R E 3 . 3 . S k e l e t o n s of d i n o s a u r f e e t : (a) Tyrannosaurus h i n d f o o t ; (b)

Iguanodon h i n d f o o t ; (c), (d) Apatosaurus h i n d a n d fore f e e t . T h e o u t l i n e u n d e r
each s k e l e t o n s h o w s t h e probable shape of its footprint.
tails of most of t h e m dragging on the ground. There is more about this

in chapter 5.
Some sauropod footprints are 1 0 - 1 5 centimeters ( 4 - 6 inches) deep.
They were probably deeper w h e n first formed because the mud m u s t
have lost water as it compacted to form rock, but they cannot have
been very m u c h deeper. If there had been too m u c h water in the mud,
it would have been too sloppy to take a footprint. T h e ground m u s t
have been fairly soft, but not sloppy.
Soft ground may have been dangerous for big dinosaurs, because of
their size. Imagine two dinosaurs of identical shape, one twice as long
as the other. It is twice as long, twice as wide and twice as high, so it
is eight times as heavy (2 = 8). T h e soles of its feet are twice as long
3
and twice as wide, so they have four t i m e s the area (2 = 4). Eight 2
times the weight has to be supported on four times the area, so the
pressure under the big dinosaur's feet is twice as m u c h as under the
small one. The big dinosaur is more likely to sink in and get bogged
down.
T h e argument seems clear, but it is a bit too simple. Suppose there
were a thick layer of soft m u d w i t h firm ground below. Small animals
DINOSAUR FOOTPRINTS 3I
might get bogged down but big ones w i t h longer legs might be able to
walk, with their feet sinking to the firm ground.
There is another complication. Different soils support loads in dif-
ferent ways. Wet clays depend m a i n l y on cohesion between the clay
particles, and the load they can support is about proportional to the
area carrying the load. Dry sand depends mainly on friction between
the grains and the load it can support is about proportional to the 1.5
power of the area: that m e a n s that four t i m e s the area can support
eight times the load. O u r big dinosaur would be more likely than the
small one to get bogged down in wet clay, but the danger of sinking
in dry sand would be about the same for both.
Real animals of different sizes are not the same shape, like these
imaginary dinosaurs. Table 3.1 shows masses and foot areas of various
animals, based on the best data I can find. Here is how the table works.
The mass of a particular Apatosaurus was probably about 35 tonnes
or 35,000 kilograms (table 2.2). Its weight (mass multiplied by gravi-
tational acceleration) was therefore 35,000 x 10 = 350,000 n e w t o n s or
350 kilonewtons. Some of the biggest k n o w n sauropod footprints are
about the right size to have been made by it. T h e area of each fore
footprint is 0.16 square meters and that of each hind print is 0.43 square
meters, giving a total (two fore and t w o hind feet) of about 1.2 square
meters. When the animal stood, this area supported 350 kilonewtons,
so (weight/area) was 290 kilonewtons per square meter. It is easy to
F I G U R E 3.4. F o o t p r i n t s a t D a v e n p o r t R a n c h , T e x a s , r e d r a w n f r o m Bird (1944).

T h e rock was formed in the early part of the C r e t a c e o u s period.
FIGURE 3.5. Rear views of (a) a lizard; (b) a bird; and (c) a mammal, and their
footprints.
calculate that 1.2' is 1.31 (use the y b u t t o n on your calculator) so

5 x
w e i g h t / ( a r e a ) ' was 270 kilonewtons per cubic meter.

s
T h e other data in the table have been calculated in the same way.
The Tyrannosaurus footprint area comes from smaller theropod foot-
prints, scaled up to m a t c h the feet of the skeleton that was used for
estimating body mass. T h e Iguanodon area comes from m u c h clearer
prints than the ones s h o w n in figure 3.6.
Look at the values of (weight/area) in table 3.1. They tell us about
the danger of getting stuck in wet clay soils. In general, we expect big-
ger values for bigger animals, as the argument about the two dinosaurs
showed. Nevertheless, the value for elephants is lower than for cattle
because elephants have relatively large feet. T h e values for Tyranno-
saurus and Iguanodon are about the same as for cattle, so these ani-
mals would have been just about as good as cattle, at crossing soft wet
clay. T h e value for the huge Apatosaurus, however, is about twice as
high as for cattle. Apatosaurus might have got bogged down on ground
that was safe for cattle.
Another possible comparison is w i t h off-road vehicles such as trac-
tors and military tanks. Various t a n k s of 3 7 - 5 1 tonnes (a little heavier
than Apatosaurus) have peak pressures of 2 0 0 - 2 7 0 kilonewtons per
square meter, under their tracks. These seem close to Apatosaurus'
DINOSAUR FOOTPRINTS 33
value of 290 kilonewtons per square m e t e r until you realize that Apa-
tosaurus would have to lift its feet in t u r n w h e n it walked. Peak forces
on the feet of people and animals during walking are generally about
double the standing values. T h e pressure under the feet of Apatosaurus
must have reached 580 kilonewtons per square meter, w h e n it walked.
It seems that the dinosaur would be less good than a tank at crossing
soft ground.
N o w look at the values for w e i g h t / j a r e a ) ' , which tell us about the
s
danger of sinking in dry sand. In this case we have to be careful about

our comparisons. Equal values m e a n equal danger of sinking only if
the animals have feet of about the same shape, and the same n u m b e r
of feet. Apatosaurus, elephants and cattle all have four, roughly cir-
cular, feet, so comparisons between t h e m seem fair. Apatosaurus has
a higher value than elephants but a m u c h lower one than cattle. It
would be m u c h less likely than a cow to get stuck in a sand dune.
Most real soils have properties between the extremes of wet clay and
dry sand.
Fossil footprints can also tell us about the speeds of the dinosaurs
that made them. They cannot tell us as certainly as if we could watch
dinosaurs running and time t h e m w i t h stopwatches, but they can give
us estimates that are probably fairly reliable.
When people walk slowly they take short strides. When they walk
faster they take longer strides and when they run they take still longer
F I G U R E 3 . 6 . ( a b o v e ) Iguanodon bernissartensis w a l k i n g w i t h i t s fore a s w e l l

a s i t s h i n d feet t o u c h i n g t h e g r o u n d a n d ( b e l o w ) f o o t p r i n t s w h i c h i t m a y h a v e
m a d e . From N o r m a n 1980. Both the s k e l e t o n and t h e footprints are from early
Cretaceous rocks.
strides (figure 3.7). Notice how stride length is defined: it is the dis-
tance from one footprint to the same point on the next print of the
same foot. Figure 3.8 shows stride lengths and speeds for h u m a n adults.
If you find a set of footprints you can measure the stride length and
use this graph to estimate how fast the person was going.
T h e same graph shows the same thing for some animals. The faster
they go, the longer their strides. Watch an adult walking with a small
child. T h e adult takes a few long strides while the child takes a lot of
short ones. Similarly, small animals take shorter strides than large ones,
at the same speed. Dogs take shorter strides than camels (figure 3.7).
Does this mean that to estimate speeds from stride lengths we need
separate graphs for each species, and separate graphs for adults and young
of each species? If that were true, dinosaur footprints could tell us
nothing about dinosaur speeds because we could never get the data for
the special graphs for dinosaurs. Fortunately, it seems not to be true.
There is a general rule that applies to birds and m a m m a l s and probably
also to dinosaurs. It seems better to compare dinosaurs to birds and
m a m m a l s than to modern reptiles, because dinosaur footprints show
that they walked w i t h their feet well in under the body (figure 3.5).
How then should we look for a general rule? What we want to do is
to m a k e a graph like figure 3.8 that will apply to animals of different
sizes. We expect long-legged animals to take long strides and short-
legged animals to take short ones, so it seems sensible to calculate
relative stride lengths
relative stride length = (stride length)/(leg length).
TABLE 3.1 Data about the danger of getting stuck in soft ground.
WEIGHT WEIGHT
MASS FOOT AREA AREA |AREA| 1 , S
Tonnes Square Meters kN/m2

KN/m"
Apatosaurus 35 1.2 290 270
Tyrannosaurus 7 0.6 120 150
Iguanodon 5 0.4 120 190
African elephant 4.5 0.6 70 90
Domestic cattle 0.6 0.04 150 740
Human 0.07 0.035 20 110
S O U R C E : D a t a are from table 2.1 and A l e x a n d e r 1985.

FIGURE 3.7. Footprints of a man walking slowly, walking fast, and running.
Leg length could be defined in various ways, but we will use the height
of the hip joint from the ground in normal standing (figure 3.9).
We may expect animals of different sizes to have equal relative stride
lengths, when running at equivalent speeds. That begs a question: what
do we mean by equivalent speeds?
The same question faces ship designers, w h o w a n t to do tests on
models before building real ships. (That way they avoid expensive mis-
takes.) They want to build ships that are cheap to run, needing little
power to propel t h e m . Much of the power for driving a ship is needed
to push along the bow wave, that builds up in front of the bow and
spreads out to either side. When a model is being tested it m u s t be
moved at the right speed to m a k e a bow wave of the same height (rel-
ative to the size of the model) as the wave that the real ship will m a k e .
This is not the same as the speed of the full-sized ship: it is an equiv-
alent speed.
Physical theory says that equivalent speeds, for this purpose, m e a n
equal dimensionless speeds, calculated in this way:
Suppose a real ship, 300 meters long, is to travel at 15 meters per sec-
ond. T h e gravitational acceleration is 10 m e t e r s / s e c o n d so the di-
2
mensionless speed is If you w a n t to m a k e tests

on a 5-meter model you should run it at 1.9 meters per second to get
the same dimensionless speed:
If that were a special rule that applied only to ships it would be no
use to us. It is actually a special case of a general rule that applies to
any motion in which gravity is important. Gravity is important in the
motion of ships because it pulls downward on the bow wave, trying to
flatten it. It is important in walking and running because of its effect
on the rise and fall of the body.
The general rule uses a concept called dynamic similarity. Imagine

you have films of two animals running, a large animal and a small one.
Show these films side by side, using t w o projectors. You could adjust
the sizes of the images, m a k i n g the animals look the same size, by
setting up the projectors at different distances from the screen. You
could adjust the t e m p o of the a n i m a l s ' running, so that both seemed
to take the same n u m b e r of strides per second, by running the projec-
tors at different speeds. You might find that the adjustments made the
two films look identical. T h a t would happen if the two animals moved
in dynamically similar fashion. D y n a m i c similarity means that the
m o v e m e n t s of one could be made identical to those of the other by
multiplying all lengths by one factor, all t i m e s by another factor, and
all forces by a third factor.
Physical theory says that if gravity is important, dynamic similarity
is only possible for a n i m a l s (or ships, or anything else) moving with
equal dimensionless speeds. Any suitable length can be used for cal-
culating dimensionless speeds. Hull length is used for ships, but leg
length is used for animals.
Animals moving w i t h equal dimensionless speeds do not have to

make dynamically similar m o v e m e n t s , but there is a good reason for
expecting t h e m to. If they are to move in the m o s t economical way
possible, so that their muscles have to do least work, they m u s t move
in dynamically similar fashion. A m o n g other things, this m e a n s that
they m u s t use equal relative stride lengths. If we plot graphs of relative
stride length against dimensionless speed, we can expect to get near-
identical graphs for similar animals of different sizes.
Figure 3.10 shows that we do. It includes all the data from figure 3.8
plus some more. It includes m a m m a l s ranging in size from dogs to an
elephant, and even a bird (the ostrich). It includes bipeds (humans and
the ostrich) as well as quadrupeds. Nevertheless, all the points lie rea-
sonably near the line. Strictly speaking, dynamic similarity is only pos-
sible between animals of the same shape. Obviously, ostriches are not
the same shape as people, dogs, or elephants, but all of these use about
the same relative stride length, at any particular dimensionless speed.
It seems likely that the same rule will also hold for dinosaurs. We can
use the graph to get rough estimates of dinosaur speeds.
Before using it, we m u s t get rid of one more worry. All the data are
for animals running on hard ground, where they did not leave visible
footprints. T h e dinosaur stride lengths that we k n o w come from foot-
prints left in ground that m u s t have been fairly soft. Does the rela-
tionship between stride length and speed that holds on hard ground
hold also on soft?
When I thought of this worry I was spending C h r i s t m a s with my
family in Norfolk, England. I took my children (then aged 11 and 13)
F I G U R E 3.9. D i a g r a m s s h o w i n g t h e m e a n i n g of leg l e n g t h .
to a nearby beach at low tide. T h e top of the beach was firm sand, and
we left only slight footprints there. Half way down the beach was sof-
ter sandy mud, and we made fairly deep footprints. At the bottom, near
the water's edge, was really horrible soft m u d and we sank in almost
to the tops of our rubber boots. At each of these levels we marked out
a 25-meter course. We walked and ran over these courses at various
speeds, in random order. We timed each other w i t h stop-watches and
counted our strides, so we were able to calculate stride lengths and
speeds. We found that we could not run fast in the soft mud—indeed,
we could hardly run in it at a l l — b u t at any speed we could manage,
softness seemed to have no effect on stride length. Each of us used the
same stride length at any particular speed, however firm or soft the
ground. This encouraged me to hope that a graph like figure 3.i0 would
give reliable estimates of dinosaur speeds, even if the dinosaurs were
on soft ground.
O n e more thing was needed, before the graph could be used. I needed
a way of estimating dinosaur leg lengths from their footprints. To find
one, I measured a lot of dinosaur skeletons. I measured the length of
the part of the foot that would rest on the ground and m a k e a footprint,
and also the height of the hip joint from the ground. (There is a danger
of error here, if the skeleton has been put together w i t h the knee un-
naturally bent or extended.) I found that leg length was about four times
foot length in a wide variety of dinosaurs, both bipeds and quadrupeds,
and decided to assume that as a general rule.
At last we are ready to estimate a speed. Take the case of the large
theropod in figure 3.1. Its foot length (measured from the m o s t com-
plete prints) was 0.64 meter, so its leg length can be estimated as 2.56
meters. Its stride length was 3.31 meters so its relative stride length
was 3.31/2.56 = 1.3. T h e graph (figure 3.10) shows that we can expect
this relative stride length when the dimensionless speed is 0.4. For an
animal of leg length 2.56 meters, that m e a n s a speed of 2.0 meters per
second:
That estimate of speed is inevitably a rough one. The scatter of points

around the line in figure 3.10 shows that we cannot expect accurate
answers. Nevertheless it seems clear that the big theropod was trav-
eling quite slowly. For h u m a n s , w i t h m u c h shorter legs, 2.0 meters per
second (4.5 miles per hour) is only a brisk walking speed.
Some other speed estimates are s h o w n in table 3.2. They have been
obtained in the same way, except that the leg length estimates for the
small Winton dinosaurs are not exactly four times footprint length: the
ratio has been estimated for each group of dinosaurs. Also, some of the
speeds have been calculated by a different formula that I published be-
fore I had collected the data for high speeds in figure 3.10. This m a k e s
little difference to the results.
Table 3.2 also shows w h e t h e r each dinosaur seems to have been
walking or running. People and m a n y birds walk to go slowly and run
to go faster. Similarly, horses, dogs, and other quadrupedal m a m m a l s
walk at low speeds and use various running gaits (trotting, galloping,
etc.) at higher speeds. There are various differences between walking
and running but the most obvious is that w h e n we walk we have each
foot on the ground for more t h a n half the t i m e so there are stages in
each stride when both feet are on the ground. In running, each foot is
on the ground for less than half the t i m e so there are stages w h e n nei-
ther foot is on the ground. Similarly for quadrupeds walking, each foot
is on the ground more than half the t i m e so there are stages w h e n both
fore feet, or both hind feet, are on the ground. For quadrupeds running,
each foot is on the ground for less t h a n half the t i m e so there are stages
when neither fore foot, or neither hind foot, is on the ground.
Watch an animal speeding up, changing from a walk to a run. In
most cases, the change is abrupt. A person or a horse, for example, is

obviously walking at one m o m e n t and obviously running at the next.
Some animals (for example, sheep) m a k e the change more gradually,
but whether the change is abrupt or gradual, it occurs at a highly pre-
dictable speed. People, and m a m m a l s in general, and birds walk when
their dimensionless speed is less than about 0.7 and run w h e n it is
more than about 0.7. T h a t rule has been used in table 3.2 to distinguish
between walking and running. If the dimensionless speed estimated
from the stride length is more than 0.7, the dinosaur is marked as run-
ning. It would be hard to be sure which the gait had been, if the es-
timated dimensionless speed were close to 0.7, but all the cases in the
table seem clear cut.
Very few footprints of running dinosaurs are known, apart from the
ones at Winton. This is not surprising. People and other fairly large
m a m m a l s walk a lot, but run only occasionally. They are particularly
unlikely to run on ground soft enough to take deep footprints, except
in emergencies.
Though running dinosaur footprints are unusual, there are some that
seem to show faster running. T h e fastest recorded are some theropod
tracks in Texas: footprints of two dinosaurs, one w i t h 29 centimeter
feet and the other w i t h 38 centimeter feet, indicate speeds of 12 meters
TABLE 3.2. Estimated speeds of dinosaurs.
ESTIMATED LEG
LENGTH ESTIMATED SPEED GAIT
Meters
Per Miles
Meters Second Per Hour
d a t a f r o m D a v e n p o r t R a n c h (fig. 3.4)
large t h e r o p o d 2.0 2.2 (4.9) walk
small theropod 1.0 3.6 (8.1) run
large sauropod 3.0 1.0 (2.2) walk
small sauropod 1.5 1.1 (2.5) walk
d a t a f r o m W i n t o n (fig. 3 . 1
large t h e r o p o d 2.6 2.0 (4.5) walk
small theropods 0.13-0.22 3.0-3.5 (6.7-7.8) run
ornithopods 0.14-1.6 4.3-4.8 19.6-10.7) am

per second (27 miles per hour). T h e larger of these, made by an animal
of (probably) about 0.6 tonne, seems also to be the largest set of di-
nosaur footprints to show running. All footprints that I know of larger
dinosaurs seem to show walking.
The top dinosaur speed, of 12 meters per second, is fast. Good hu-
man athletes sprint at up to 10 meters per second, racehorses at up to
17 meters per second, and greyhounds at up to 16 meters per second.
These speeds are known accurately, because they are measured in races.
Remarkably little is k n o w n about top speeds of other animals. With
colleagues, I have measured the speeds of various African animals by
filming t h e m from a vehicle, while chasing t h e m across country on
the grassy plains of Kenya. Zebras, giraffes, and various antelopes gal-
loped at top speeds of 11 to 14 m e t e r s per second. Ostriches seemed
to be a little faster. It seems possible that the racers bred by man, race-
horses and greyhounds, are the fastest land animals. This would not
be surprising. They have been bred for speed alone, but the evolution
of wild animals has been guided by other needs, as well as speed.
Many books give higher top speeds for wild animals, up to an amaz-
ing (incredible?) m a x i m u m of 30 meters per second for cheetahs. N o n e
of the books that give these speeds explain in detail how they were
measured. There are m a n y possible sources of error. For example, imag-
ine you are driving in a vehicle alongside an animal, watching your
speedometer. It swerves away from you, and you swerve to keep up
with it. You are now traveling on the outside of a bend, w i t h it on the
inside. You m u s t drive faster than it is running, to keep beside it. Er-
rors of this kind were avoided in the experiments that gave top speeds
of 11 to 14 meters per second. T h e m e t h o d s used were explained in
detail in a scientific journal, so that other scientists could criticise t h e m
if they were faulty.
That means that the estimated top speed of 12 meters per second,
for the Texas theropods, would probably be a very creditable speed for
an antelope. Those dinosaurs were not particularly large, but they were
fast.
We will ask one more question of the footprints. T h e Winton foot-
prints (figure 3.1) and the Davenport Ranch ones (figure 3.4) were made,
in each case, by m a n y individual dinosaurs. Were these dinosaurs mov-
ing around in groups and, if so, w h a t were they doing?
In both cases they are suspected of moving in groups, but the evi-
dence is inconclusive. In both cases there are a lot of tracks running
in the same direction, which suggests group m o v e m e n t . All the prints
seem to have been fresh w h e n silted over and therefore (probably) were
made about the same time. There is no indication of some older prints
being more weathered by rain or other causes than later prints.
At Davenport Ranch there are tracks of 23 sauropods, all walking in

the same direction. They all had similar-shaped feet and probably be-
longed to the same species, but some had feet twice as large as others.
T h e largest were probably eight times as heavy as the smallest but all
the sauropods, large or small, seem to have been walking at about the
same low speed. T h e impression is of a herd, adults and juveniles, trav-
eling together at a leisurely pace.
At Winton the picture seems more exciting. T h e single large ther-
opod seems to have been walking southwest. More than 150 small di-
nosaurs, theropods and ornithopods, ran m u c h faster in the opposite
direction, toward the northeast. Richard Thulborn and Mary Wade, who
studied the tracks, tell this story. T h e area was a m u d bank at the edge
of a lake. A large group of small dinosaurs were on the bank where
they had come to drink or to h u n t for food. The ornithopods may have
been feeding on plants and the small theropods may have been trav-
eling w i t h them, ready to catch and eat any insects that they disturbed.
Both the ornithopods and the theropods varied in size, but some of
t h e m were probably juveniles. T h e small theropods may all have been
one species and all but one of the ornithopods m a y have been another
species. (The exception had much larger feet than any of the other or-
nithopods.)
Suddenly the big theropod arrived, walking off the mainland onto
the m u d b a n k . It was m u c h bigger t h a n any of the other animals, well
able to attack and eat any of t h e m . T h e small dinosaurs were threat-
ened, but the way forward was blocked by the water. All they could
do was to double back past the theropod onto the mainland, running
as fast as they could on the soft ground.
This chapter has told about fossil footprints of dinosaurs and has
shown how the speeds of animals that made t h e m have been esti-
mated, from the spacing of the prints. Most of the speeds indicated by
dinosaur footprints (especially by big ones) are slow, but a few foot-
prints of running dinosaurs are known. T w o exceptional trackways seem
to have been made by dinosaurs running at speeds that would be fast
for antelopes.
I also discussed w h e t h e r big dinosaurs would be in danger of sinking
in soft ground. Apatosaurus would have been more apt to sink than a
cow in soft clay, but would be less likely than a cow to sink in a sand
dune.
Principal Sources
T h e D a v e n p o r t R a n c h a n d W i n t o n t r a c k s h a v e b e e n d e s c r i b e d b y Bird ( 1 9 4 4 | and
T h u l b o r n a n d W a d e (1984), r e s p e c t i v e l y . N o r m a n (1980) d i s c u s s e d t h e s u p p o s e d
Iguanodon t r a c k . F a r l o w (1981) d e s c r i b e d t h e very fast t h e r o p o d t r a c k .
DINOSAUR FOOTPRINTS 4.?
Alexander (1985) collected t h e data a b o u t pressures on t h e ground. A l e x a n d e r (1976)

i n t r o d u c e d t h e m e t h o d o f e s t i m a t i n g s p e e d s from s t r i d e l e n g t h s . T h e " F r o u d e n u m -
b e r " defined i n t h a t p a p e r i s t h e s q u a r e o f t h e q u a n t i t y called " d i m e n s i o n l e s s s p e e d "
i n t h i s b o o k . T h e d a t a o n s t r i d e l e n g t h s o f m o d e r n a n i m a l s c o m e s largely from Alex-
a n d e r and Jayes (1983) a n d t h e d a t a o n m a x i m u m s p e e d s from A l e x a n d e r , L a n g m a n ,
and Jayes (1977).
A l e x a n d e r , R. M c N . 1976. E s t i m a t e s of s p e e d s a n d d i n o s a u r s . Nature 2 6 1 : 1 2 9 - 3 0 .
Zoological journal of the Linnean Society 8 3 : 1 - 2 5 .
A l e x a n d e r , R. M c N . a n d A. S. Jayes. 1983. A d y n a m i c s i m i l a r i t y h y p o t h e s i s for t h e
g a i t s of q u a d r u p e d a l m a m m a l s . Journal of Zoology 2 0 1 : 1 3 5 - 1 5 2 .
A l e x a n d e r , R. M c N . , V. A. L a n g m a n , a n d A. S. Jayes. 1977. Fast l o c o m o t i o n of s o m e
African u n g u l a t e s , lournal of Zoology 1 8 3 : 2 9 1 - 3 0 0 .
Bird, R. T. 1944. D i d b r o n t o s a u r u s e v e r w a l k on land? Natural History, New York
53:60-67.
Farlow, J. O. 1 9 8 1 . E s t i m a t e s of d i n o s a u r s p e e d s from a n e w t r a c k w a y s i t e in T e x a s .
Nature 2 9 4 : 7 4 7 - 7 4 8 .
N o r m a n , D. B. 1980. On t h e o r n i t h i s c h i a n d i n o s a u r Iguanodon bernissartensis of
Bernissart (Belgium). Memoires de l'lnslitut Royal des Sciences Nalurelles de Bel-
gique 1 7 8 : 1 - 1 0 3 .
T h u l b o r n , R . A . a n d M . W a d e . 1984. D i n o s a u r t r a c k w a y s i n t h e W i n t o n for-
m a t i o n ( m i d - C r e t a c e o u s ) of Q u e e n s l a n d . Memoirs of the Queensland Museum
21:413-517.
IV
Dinosaur Strengths
F seem to tell us that large dinosaurs usually moved

OSSIL F O O T P R I N T S
slowly, but could they move fast in an emergency? Could they

jump, and do other athletic things? We expect very large animals to be
lumbering monsters. Buffaloes are less nimble than gazelles and ele-
phants are less n i m b l e than buffaloes. Were dinosaurs that were m u c h
larger than elephants less nimble still?
You have to be strong to run fast or to jump. When you stand, your
weight is divided between your two feet: the force on each foot equals
half your weight. When you walk, for m u c h of the t i m e you have only
one foot on the ground and the peak force on each foot is about equal
to body weight. When you run, each foot is on the ground for less time
so the peak forces have to be bigger. T h e average vertical force, over
a complete stride, m u s t equal body weight, so if the feet are on the
ground for less of the t i m e the peak forces m u s t be bigger. In jogging,
each foot is on the ground for about 35 percent of the time and the
peak force is about 2.7 t i m e s body weight. In sprinting, each is on the
ground for about 22 percent of the t i m e and the peak force is about 3.5
times body weight. These forces have been measured by means of force
plates, force-sensitive panels set into the ground. The faster you run,
the bigger are the forces, and even bigger forces may act when you
jump.
Imagine t w o animals of exactly the same shape, one twice as long
as the other. It is twice as long, twice as wide, and twice as high, so
it is eight times as heavy. Its bones and muscles have twice the di-
ameter, and four t i m e s the cross-sectional area. Suppose that these two
animals perform the same activity, moving in dynamically similar
fashion. All the forces involved are in proportion to their weight, eight
times as large for the larger animal. T h e strengths of their bones and
muscles are proportional to their cross-sectional areas, only four times
DINOSAUR STRENGTHS 4.5
as large for the larger animal. This m e a n s that the big animal is work-
ing nearer the limit than the small one, the limit set by the strengths
of its bones and muscles. If the two animals try something more stren-
uous, the small animal may succeed but the large one may not.
That argument does not apply directly to real animals, because an-
imals of different sizes are not the same shape, but it helps us to un-
derstand why buffaloes are less athletic than gazelles. It also m a k e s it
clear that athletic ability depends on the strengths of bones and m u s -
cles.
We do not know exactly h o w big dinosaur muscles were, because
they are not preserved in fossils, so we cannot easily estimate their
strengths. We have plenty of dinosaur bones but we cannot usefully
measure their strengths, because fossil bones are not made of the same
stuff as living ones. Bone consists largely of fibers of the protein col-
lagen reinforced by crystals of the mineral hydroxyapatite. In fossils
the protein has decayed and the bone m a y have been impregnated by
other materials seeping through the rock.
Though the composition of bones is changed by the processes of fos-
silization, their sizes are probably not. We can use the dimensions of
the fossil bones to estimate the strengths of the bones in the living
dinosaurs, if we assume that dinosaur bone was as strong as bone from
modern animals. T h a t assumption seems plausible. Slices of dinosaur
bone, examined under a microscope, look very like slices of bone from
modern m a m m a l s . Samples of bone from the shafts of leg bones of
various birds and m a m m a l s all have similar properties. (I cannot find
any records of strength tests on reptile bone.)
We need to know more about stresses and strains. These words are
used in rather vague ways by most people, but are given precise mean-
ings by engineers. Figure 4.1a represents a bar of some material; it does
not matter whether it is bone, plastic, rubber or s o m e t h i n g else. One
of its ends is firmly fixed by being embedded in a rigid wall. T h e grid
of squares will help us to see how the bar is deformed w h e n forces act
on it.
In figure 4.1b a force is pulling on the free end of the bar, stretching
it. The strain is the relative change of length:
change of length
strain = — —
initial length
Obviously, more force is needed to produce a given strain in a thick

bar than in a thin one of the same material. Also more force is needed
to break a thick bar than a thin one. T h i s m a k e s it useful to calculate
the stress:
46 DINOSAUR STRENGTHS
force
stress =
cross-sectional area
If the stress reaches a certain limit, called the tensile strength of the
material, the bar will break.
In figure 4.1c a force is pushing on the free end of the bar, com-
pressing it. Strain and stress are defined in the same way as before but
in this case the strain is negative, because the bar has been made shorter.
Also, by convention, the stress is regarded as negative. If the stress
reaches a limit, called the compressive strength, the bar will break.
T h e tensile strength of m a m m a l bone is about 160 n e w t o n s per square
millimeter and the compressive strength is about - 2 7 0 newtons per
square millimeter.
In figures 4.1b and c the forces are in line with the bar but in figure
4.Id a force acts at right angles to the bar, bending it. Notice how the
bar is stretched on the outside of the bend and compressed on the in-
side. T h e strain and stress are positive on the outside of the bend, and
negative on the inside. If the stress on the outside reaches the tensile
F I G U R E 4 . 1 . A bar fixed rigidly at o n e end, a c t e d on by v a r i o u s forces. T h e s e

d i a g r a m s are explained in t h e text.
DINOSAUR STRENGTHS 47
strength, or if the stress on the inside reaches the compressive strength,

the bar will break.
In figures 4.1b and c the stresses are the same all along the bar, but
in figure 4.Id they are not. T h e biggest stresses are at the fixed end of
the bar, where the force has m o s t leverage. If the force is increased
until the bar breaks, it will break where it meets the wall.
Consider one particular cross section of the bar, at XX. This section
is at a distance x from the line of the force F. T h e bending effect of
the force at this particular section is described by the bending m o m e n t
Fx (the force multiplied by the distance). T h e stresses in this section
range from Fx/Z at the top surface of the bar (on the outside of the
bend) to -Fx/Z at the b o t t o m surface (on the inside of the bend). In
these formulae, Z is a quantity called the section modulus, that de-
pends on the size and shape of the cross section and also on the di-
rection of bending. Its value for a circular cross section is 0.78 t i m e s
the cube of the radius. Engineering textbooks explain how it can be
calculated for other shapes, including irregular ones.
A big section m o d u l u s m a k e s a bar hard to break. Figure 4.2 shows
some sections that have equal cross-sectional areas, but different sec-
tion moduli. T h e tube and the I beam have larger section moduli than
the cylindrical rod. Bicycle frames are made of tubes, and girders are
given I sections, to m a k e t h e m stronger against bending m o m e n t s t h a n
simple rods would be.
N o w we are ready to start thinking about stresses in dinosaur leg
bones. Figure 4.3a represents the leg skeleton of a r u n n i n g dinosaur.
The foot is pressing down on the ground and the ground is pressing up
on the foot. The precise direction of the force m a y vary, depending on
the stage of the stride and on w h e t h e r the animal is accelerating or
decelerating, but we will suppose that the direction is as shown by the
arrow. The foot is pivoted at the ankle, so this force tries to rotate the
foot counterclockwise about the ankle joint. A balancing force is needed,
trying to rotate the foot clockwise. This would be supplied by calf m u s -
cles which would probably be arranged as s h o w n in the diagram. (Birds
and crocodiles have muscles like this.)
We m u s t t h i n k about the balance of forces in m o r e detail. In figure
4.3b two forces are s h o w n acting on the sole of the foot instead of one:
a force P parallel to the tibia and a force R at right angles to it. These
two forces acting together would have exactly the same effect as the
single force shown in figure 4.3a, but it will be convenient to break
the force down into these c o m p o n e n t s : it will help us to t h i n k about
the stresses in the tibia.
In figure 4.3b, the ground exerts forces P and R on the sole of the
foot, and the calf muscles exert force M on the foot behind the ankle.
FIGURE 4 . 2 . Possible shapes for the cross section of a beam: (a) is a solid rod,
(b) a tube and (c) an I-beam. All have the same cross-sectional area. The num-
bers are section moduli for bending by vertical forces, relative to the value for
the solid rod.
The only other forces on the foot act at the ankle joint, where the tibia
presses against the ankle bones. Notice that all the forces shown in
figure 4.3b are either parallel to the tibia or at right angles to it. T h e
forces in each of these directions m u s t balance. Forces M and P act
upward, parallel to the tibia, so the tibia m u s t press downward with a
balancing force M + P. Force R presses backward on the sole of the
foot so the tibia m u s t press forward at the ankle, with a force R.
N o w think about the forces on the lower end of the tibia. The tibia
presses downward on the foot w i t h a force M + P so the foot m u s t
press upward on the tibia with equal force (figure 4.3c). The tibia presses
forward on the foot with a force R so the foot presses backward on the
tibia w i t h an equal force. Force M + P, acting along the shaft of the
tibia, compresses the bone but force R acting at right angles to the shaft
of the tibia, bends it.
A cross section of the tibia is picked out by shading in figure 4.3c.
This section has area A and section m o d u l u s Z, and is at a distance x
from the end of the bone. Force [M + P), acting alone, would set up a
stress - [M + P)/A in the cross section (remember that compressive
stresses are negative). Force R, acting alone, would exert a bending mo-
m e n t Rx and set up stresses ranging from Rx/Z at one edge of the
section to -Rx/Z at the opposite edge. T h e stresses produced by the
two forces acting together can be calculated by adding up their separate
effects. T h u s the total stress ranges from — (M + P)/A + Rx/Z at one
edge of the section to - (M + P)/A - Rx/Z at the other.
This analysis tells us that the forces at the ankle would have both
a compressing effect on the tibia and a bending effect. It also shows
how the stresses could be calculated if the forces on the feet were known.
The tibia has been used as an example but similar calculations could
be made for the other long bones of the legs.
This could be the starting point for a fearsomely elaborate series of
calculations. We could try to reconstruct the precise sequence of move-

ments made by running dinosaurs, perhaps by making animated car-
toons. We could calculate the forces that would act on the feet and use
these, together w i t h the bone angles taken from the cartoons, to cal-
culate the forces M + P and R on each of the major leg bones. Then,
if we knew the dimensions of the bones, we could calculate the stresses
in them. We could do this for several different running speeds, calcu-
lating the stresses for each. We would find that higher speeds produced
bigger stresses and we would be able to e s t i m a t e the speeds at which
the bones would break if the muscles could m a k e the animals run so
fast. M a x i m u m running speeds would be less t h a n this, allowing some
margin of safety.
Such calculations would raise all sorts of doubts. Did dinosaurs take
long strides or short ones? H o w long did the foot remain on the ground,
in each stride? Did they run w i t h their legs relatively straight like el-
ephants or bent like smaller animals? Did they get the foot into a par-
ticular position by bending the knee while keeping the ankle relatively
straight or by bending the ankle while keeping the knee straight? Were
bone stresses at m a x i m u m running speeds almost enough to break the
bones, or was there a wide margin of safety?
F I G U R E 4 . 3 . F o r c e s o n t h e foot a n d t i b i a o f Tyrannosaurus. D i a g r a m (a) s h o w s

t h e leg s k e l e t o n , t h e force o n t h e sole o f t h e foot a n d t h e m u s c l e s t h a t e x t e n d
t h e a n k l e ; (b) s h o w s all t h e i m p o r t a n t f o r c e s o n t h e foot; a n d (c) s h o w s f o r c e s
on the lower end of the tibia.
I faced all these questions in my research on moas, the giant ostrich-

like birds that are discussed in chapter 11. Here we avoid facing t h e m
directly by m a k i n g the general assumption, that the m o v e m e n t s of di-
nosaurs and the stresses developed in their skeletons were m u c h the
same as for modern animals. We also avoid complicated calculations.
I believe that the simple m e t h o d I am going to describe gives at least
as good an indication of the athletic ability of dinosaurs as any other
m e t h o d so far invented.
I m u s t m a k e my assumptions clear. T h e first assumption is that di-
nosaurs moved m u c h like modern animals. More precisely, I will as-
s u m e that the m o v e m e n t s of dinosaurs were dynamically similar to
those of modern m a m m a l s , traveling at the same dimensionless speed.
I explained w h y this assumption seems reasonable, when I discussed
dinosaur footprints (chapter 3). Precise dynamic similarity is impos-
sible unless the animals being compared are exactly the same shape,
but I will m a k e the best comparisons I can. I will compare sauropods
w i t h elephants, which are obviously not the same shape: sauropods
have relatively small heads on long necks and elephants have large heads
on short necks. Nevertheless, the proportions of the legs are similar
(figure 4.4). Both sauropods and elephants have relatively long femurs
and short toes, and it seems likely that sauropods moved their legs in
m u c h the same way as elephants. However, elephants have big heads
and puny tails, so m o s t of their weight is supported by their front legs.
Sauropods like Apatosaurus had huge hindquarters and tails, so most
of their weight m u s t have been supported by their hind legs. I will
allow for this difference.
T h e second assumption is about stresses. I will assume that the
thicknesses of dinosaur leg bones were adjusted so that, when the di-
nosaur ran as fast as it could, the stresses in its leg bones were the
same as in the leg bones of modern animals, running as fast as they
can. For example, I have calculated that the peak tensile and com-
F I G U R E 4 . 4 . O u t l i n e s o f Apatosaurus a n d a n A f r i c a n e l e p h a n t , w i t h s o m e o f
t h e b o n e s d r a w n in. T h e o u t l i n e of t h e e l e p h a n t h a s b e e n traced from a film
o f fast r u n n i n g . A l e x a n d e r e t al. 1 9 7 9 .
DINOSAUR STRENGTHS 5 I
pressive stresses in the h u m e r u s of an elephant running fast were about

+69 and - 8 5 n e w t o n s per square m i l l i m e t e r (about one third of the
stresses that would break the bone). I will assume that w h e n sauropods
ran as fast as possible the stresses in their bones were about the same.
This implies that, if dinosaur bone had the same strength as elephant
bone (as is likely), sauropod leg bones had the same margin of safety
as elephant leg bones. This assumption will help us to decide w h e t h e r
sauropods were probably more athletic than elephants, or less.
Our calculations would still be complicated, were it not for some-
thing very convenient. We can ignore the forces parallel to the bone
(M and P in figure 4.3) and consider only the force at right angles to it
[R). This may seem odd, because (M + P) is usually m u c h larger than
R. (Leg muscles attach close to the joints, so their lever arms are short
and the forces M that they have to exert are usually m u c h larger than
the forces on the feet.) However, forces at right angles to leg bones are
much more effective at producing stresses than forces acting along their
lengths. This is because the bones are long and relatively thin. It is
m u c h easier to break a bone, or a stick, or any other long thin object,
by bending it than by compressing it lengthwise.
The most direct evidence we have about the relative importance of
compression and bending, in the leg bones of animals, comes from ex-
periments in which strains have been measured in bones of living an-
imals. These experiments used strain gauges, small devices made of
metal foil m o u n t e d on plastic. Stretching or compressing a strain gauge
changes its electrical resistance, so the gauge can be used to measure
strain at the surfaces of structures. In surgical operations on horses,
sheep, and other animals, strain gauges were glued to the surfaces of
leg bones. The skin was sewn up again, but wires from the strain gauges
were brought out through the incision so that they could be connected
to recording equipment whenever strain was to be measured. Animals
recover quickly from the operation and can soon run normally. T h e
operation is so simple and painless that Dr. Hampson, a m e m b e r of a
team doing this kind of experiment, allowed his colleagues to perform
the operation on him, attaching a strain gauge to his tibia.
Figure 4.5 shows records from a jumping pony. It had two strain gauges
glued opposite each other on a leg bone, one on the front surface of the
bone and one on the rear. T h e records show strains indicating positive
(tensile) stresses up to 50 newtons per square millimeter under the front
gauge during landing and negative (compressive) stresses up to 60 of
the same units under the rear one. T h e positive stresses almost m a t c h
the negative ones, indicating that most of the stress was due to bending
(as in figure 4.Id) rather than lengthwise compression (as in figure 4.1c).
Experiments on various leg bones of horses, sheep and dogs gave sim-
FIGURE 4.5. Stresses calculated from records of strain in the front and rear
surfaces of a leg bone (the radius) of a pony, which took off for a jump at "off"
and landed at "on." From Bicwener, Thomason, and Lanyon 1983.
ilar results in nearly every case. There seems to be a general rule that
bending m o m e n t s are usually m u c h more important than lengthwise
compression, in causing stresses in bones.
T h i n k about the example of the elephant h u m e r u s . The calculated
stresses at the surfaces of the bone were + 6 9 and - 8 5 newtons per
square millimeter. They were made up of -8 n e w t o n s per square mil-
limeter, in each case, due to lengthwise compression and + 77 or - 7 7
n e w t o n s per square m i l l i m e t e r due to the bending m o m e n t . ( - 8 + 77
= 69 and -8 - 77 = - 8 5 . ) If I had simplified my calculations by con-
sidering only the bending m o m e n t I would have obtained +77 and - 7 7
instead of - 6 9 and - 8 5 , and would not have been far wrong. We will
consider only stresses due to bending m o m e n t s .
T h e stresses due to bending m o m e n t s in a cross section of a bone
range from +Rx/Z to -Rx/Z (figure 4.3). Suppose that we measure the
distance x and the section m o d u l u s Z for corresponding sections of the
same bone, in t w o animals. If the animals m a k e dynamically similar
m o v e m e n t s , their forces R will be in proportion to the weight W that
the legs have to support. (If the animals are bipeds, W are the weights
of their bodies. If they are quadrupeds, W m e a n s the part of the body
weight supported by the fore or hind legs, whichever are being consid-
ered.) T h u s the stresses due to bending m o m e n t s will be proportional
to Wx/Z. T h e stresses in the two animals will be almost equal, when
they m a k e dynamically similar m o v e m e n t s , if they have equal values
of Wx/Z.
This suggests that we can use Z/Wx (which is Wx/Z turned upside
down) as an indicator of athletic ability. T h e bigger Z/Wx is, the less
stress will act, in any particular activity. Large values of Z/Wx mean
that strenuous activities are possible w i t h o u t setting up dangerous
stresses in the bones. We can estimate the athletic prowess of dino-
saurs by calculating values of Z/Wx and comparing t h e m w i t h values
for modern animals, whose athletic ability we know. Whichever ani-
mal has the larger values of Z/Wx is likely to be the more athletic.
We are going to apply this formula to quadrupedal dinosaurs, so we
need to know how to measure W, the weight supported by the fore or
hind legs. The m e a s u r e m e n t is easily made for living animals, if a force
plate is available. When the animal stands w i t h all four feet on the
plate, the vertical force registered by the plate is the total weight of
the animal. When it stands w i t h its fore feet on the plate and its hind
feet on the floor alongside, the force registered is the weight supported
by the fore feet. If the animal is too heavy for an ordinary force plate,
a weighbridge of the kind used for weighing trucks can be used instead.
Measurements on a weighbridge showed that an elephant carried 58
percent of its weight on its front legs and 42 percent on its hind legs.
We cannot get living dinosaurs to put on weighbridges so we have
to use models, and there is a problem about doing that. T h e models I
used were solid plastic, of the same density throughout, but different
parts of the living dinosaur had different densities. Bone is about twice
as dense as flesh and guts (i.e. the same volume is about twice as heavy).
If a dinosaur had a lot of bone at its front end and very little in its tail
(for example) its fore feet would have to support a bigger fraction of
the total weight than they do in a plastic model. T h e bones of dino-
saurs were not concentrated near one end, but distributed all through
the body, so we probably do not need to worry about the model not
having dense bones. Also if the front half of the dinosaur were filled
with air, the fraction of body weight carried by the fore feet would be
less than for a solid model. There m u s t have been quite a lot of air in
the lungs, which would have been in the front half of the body, and it
seemed necessary to take account of it. I therefore bored holes through
the models where the lungs would have been. Experiments on caimans
and on large m a m m a l s such as elands and camels had shown that their
lungs occupy about 8 percent of the v o l u m e of the body. I assumed
that the same would be true of dinosaurs and adjusted the sizes of the
holes so as to reduce the mass of each model by 8 percent.
Figure 4.6 shows how I did the experiment, using a letter balance
instead of a weighbridge. When I arranged the dinosaur model as shown,
the balance read 51 grams. I removed the model leaving the rubber pad
on the pan, and the reading fell to 5 grams. T h e m a s s of the model
5 4 DINOSAUR STRENGTHS
F I G U R E 4.6. An e x p e r i m e n t to discover h o w m u c h of t h e weight of a d i n o s a u r

w a s s u p p o r t e d b y i t s fore l e g s . T h e c i r c l e o n t h e m o d e l ' s c h e s t i s a h o l e b o r e d
to represent the lungs.
was 85 grams. Therefore, the fraction of the model's weight supported

by the fore legs was (51 - 5)/85 = 0.54.
T h e pads of foam rubber had an important function. The model was
rigid, so only three of its feet would probably rest on the ground when
it stood on a rigid surface. Similarly a table on a flat floor rests on just
three of its four feet unless the lengths of its legs are very precisely
matched. The pieces of foam rubber ensured that the weight was shared
between all four feet.
T h e results of the experiments are s h o w n in table 4.1. It seems that
Brachiosaurus supported almost half its weight on its fore feet, but
that the other sauropods, Diplodocus and Apatosaurus, supported most
of their weight on their hind feet. (The Apatosaurus values come from
a h o m e m a d e model and are probably less accurate than the rest.) Ste-
gosaurus also carried most of its weight on its hind feet but Triceratops
carried more weight on its fore feet.
T h e living animals in the table are not the ones we would most like
to know about. T h e elephant is an Indian one, but my m e a s u r e m e n t s
of elephant bones are from an African elephant. I will assume that the
percentages of body weight carried by the fore and hind feet are the
same for both species. I will also assume that the percentages for buf-
falo are about the same as for a horse.
At last we are ready to calculate Z/Wx, the indicator of athletic abil-

ity. My values for the section m o d u l u s Z and the distance x come from
m e a s u r e m e n t s of bones, both from modern animals and from dinosaur
fossils. The sections that I measured were usually about half way along
the bones. I k n e w the masses of the modern animals because the bones
came from weighed carcasses, and I had calculated the masses of the
dinosaurs from the volumes of models (table 2.2). I took the fractions
of body weight carried by the fore and hind feet from table 4.1, and so
was able to calculate the weight W carried by each pair of feet.
Notice that I use dinosaur masses calculated from the volumes of
models, that is masses taken from the " C o l b e r t " and "Alexander" col-
u m n s in table 2.2. It would have been wrong to use the " A n d e r s o n "
masses for this job because they were calculated from bone dimen-
sions. They assume a particular relationship between body mass and
the dimensions (and so the strengths) of the bones. If we used t h e m to
assess the athletic ability of dinosaurs, by comparing the strengths of
bones to the loads they have to carry, our argument would be circular.
The resulting values of Z/Wx are s h o w n in table 4.2. Notice that
the values for elephant bones are all m u c h lower than the values for
corresponding bones of buffalo. T h e comparison is not a particularly
good one, because the relative lengths of the leg bones of elephants and
buffaloes are very different, so their m o v e m e n t s can never be precisely
dynamically similar. However, the values do indicate correctly that el-
ephants are less athletic than buffaloes. Most m a m m a l s walk at low
TABLE 4 . 1 . Percentages of body weight supported by the fore and

hind feet of various quadrupeds.
PERCENTAGES OF BODY WEIGHT
S U P P O R T E D BY
Fore Feet Hind Feet
Indian elephant 58 42
Horse 59 41
Diplodocus 22 78
Apatosaurus 30 70
Brachiosaurus 48 52
Triceratops 54 46
Stegosaurus 18 82
speeds, trot at intermediate speeds, and gallop at high speeds. Ele-

phants generally do not trot but use an alternative running gait, the
amble, w i t h a different sequence of foot m o v e m e n t s (figure 4.4). Their
fastest gait is a slow amble, and they seem unable to gallop. Buffaloes
can gallop, though not very fast. Elephants do not jump and are often
kept in their enclosures in zoos by a narrow ditch instead of a fence.
I have no data on buffalo jumping but I have film of an impressive jump
by an eland (an antelope of about the same mass).
The quadrupedal dinosaurs in table 4.2 had legs with elephant-like
proportions, so comparisons w i t h elephants should be revealing. The
value of Z/Wx for Diplodocus femur is even lower than for the ele-
phant, which suggests that Diplodocus was even less athletic than el-
ephants: it could walk but probably could not run. The values for the
h u m e r u s , femur, and tibia of Apatosaurus are all close to the values
for elephant, which suggests that Apatosaurus was about as athletic as
an elephant, able to run but not to gallop or to jump. The values for
Triceratops are close to those for buffalo, suggesting that Triceratops
may have been able to gallop.
T h e largest modern galloping animal seems to be the White rhinoc-
eros, which is said to reach 3 tonnes, about half as m u c h as a really
large African elephant. I have drawn the rhinos in figure 4.7 from a
film of a rhinoceros galloping. It was being chased by a vehicle across
TABLE 4.2. Values of Z/Wx (the quantity used as an indicator of

athletic ability) for leg bones of dinosaurs and modern animals.
Body Mass Values of Z/Wx

(Tonnes) (Square Meters per Giganewton)"
Femur Tibia Humerus
African elephant 2.5 7 9 11
Buffalo 0.5 11 21
Diplodocus 12-19 3-5
Apatosaurus 34 9 6 14
Triceratops 6-9 15-21 12-20
Ostrich 0.04 44 18
Man 0.06 15 15
Tyrannosaurus 8 9
"A g i g a n e w t o n is o n e billion |10"| n e w t o n s .

F I G U R E 4.7. O u t l i n e s traced from a film of a W h i t e r h i n o c e r o s galloping, w i t h

d r a w i n g s o f Triceratops g a l l o p i n g b a s e d o n t h e s a m e f i l m .
a very large zoo enclosure, and reached a m a x i m u m speed of 7.5 meters

per second (17 miles per hour). If Triceratops could gallop, it presum-
ably galloped like a rhino.
There are no really large modern bipeds to compare w i t h the large
bipedal dinosaurs. T h e largest are ostriches and h u m a n s , and I have
put data for both in table 4.2. The m a n whose bones I measured had
been ill and inactive for a few years before he died so his bones may
have become thinner and weaker than w h e n he was healthy. Even so,
Z/Wx for his femur was higher than for Tyrannosaurus, but the value
for ostrich femur was much higher. These comparisons suggest (un-
surprizingly) that Tyrannosaurus was less athletic than ostriches and
people. T h e Tyrannosaurus value is close to the elephant one, which
again suggests that Tyrannosaurus could not have run very fast. T h e
proportions of Tyrannosaurus' hind legs are unlike those of ostriches,
people or elephants, so none of these comparisons are really satisfac-
tory, but they seem to be the best possible.
Whether they were strong enough for fast running or not, dinosaur
skeletons m u s t have been strong enough to support the animals w h e n
they copulated. T h e male rhino in figure 4.8a seems to be imposing a
terrific load on the female's back. Male elephants also m o u n t females
to copulate like this, but dinosaurs m u s t have done it rather differently
because their big tails would get in the way. Reptiles do not have the
F I G U R E 4.8. (a] B l a c k r h i n o c e r o s c o p u l a t i n g , f r o m a p h o t o g r a p h ; (b) Anolis l i z -

ards c o p u l a t i n g on a b r a n c h of a tree, from G r e e n b e r g a n d N o b l e 1944.
sexual orifice separate from the anus, like h u m a n s and other mam-
mals, but have the t w o combined to form a single opening called the
cloaca. This is on the underside of the tail, a short distance behind the
legs. To copulate, the male m u s t press his cloaca against the female's.
T h a t m a y sound impossible to do from behind, but lizards manage by
twisting their tails together (figure 4.8b). Dr. Beverley Halstead, a Brit-
ish paleontologist, has suggested that dinosaurs also may have copu-
lated like this, w i t h the male lifting one hind leg and putting it over
the female's back.
If they did, bigger loads would act on their hind legs than in normal
standing. When the male lifted one hind leg, the other would have to
carry twice the standing load. If he rested his leg on the female's back,
her hind legs would have to carry an increased load. However, the loads
would be no bigger than in walking, when peak forces on the feet were
probably about twice as m u c h as in standing. (I explained at the be-
ginning of this chapter why peak forces on h u m a n feet are twice as
m u c h in walking as in standing.) If dinosaurs were strong enough to
walk they were also strong enough to copulate. They were presumably
strong enough to do both.
This chapter has told a complicated story. Here, briefly, is how it
went. T h e faster an animal runs or the more athletically it behaves the
bigger, in general, are the forces on its legs. It is harder for large animals
than for small ones to withstand these forces because doubling the length
of an animal w i t h o u t changing its proportions m a k e s it eight t i m e s

heavier but only four times stronger. This is w h y elephants are less
athletic than gazelles.
I used the dimensions of dinosaur bones to assess athletic ability. I
assumed that the running movements of dinosaurs were m u c h like those
of modern animals and that their skeletons were proportioned so that
the stresses in them, in the m o s t strenuous activities, were the same
as for modern animals. I s h o w e d that, for rough calculations of stress,
only forces that set up bending m o m e n t s in t h e bones need be consid-
ered. This led to the definition of the quantity Z/Wx, that takes ac-
count of the dimensions of a leg bone and of the weight supported by
the legs. T h e larger its v a l u e s of Z/Wx, the m o r e athletic an animal is
likely to have been.
I compared quadrupedal d i n o s a u r s w i t h elephants. T h e values of Z/
Wx suggested that Apatosaurus was about as athletic as an elephant,
able to run slowly but not to gallop. Diplodocus was less athletic but
Triceratops seems to have been more athletic, and m a y have galloped
like a rhinoceros. Tyrannosaurus was m u c h less athletic than ostriches
or people, the largest modern bipeds.
Finally (and rather obviously), dinosaurs were strong enough to cop-
ulate.
Principal Sources
I have based this chapter on Alexander (1985) but have used a new method to es-
timate the weight supported by the fore and hind feet of dinosaurs. Consequently,
some of\the numbers given in the chapter are slightly different from those in the
original paper. The new method is not necessarily better than the old one but it is
more direct, and easier to explain. The information about elephants and buffaloes
comes from Alexander et al. (1979). The data from strain gauges glued to bones of
living animals are from Biewener, Thomason and Lanyon (1983).
A l e x a n d e r , R . M c N . 1 9 8 5 \ M e c h a n i c s o f p o s t u r e a n d gait o f s o m e large d i n o s a u r s .
Zoological Journal of the Linnean Society 8 3 : 1 - 2 5 .
A l e x a n d e r , R. M c N . , G. M.O. Maloiy, B. H u n t e r , A. S. Jayes, a n d J. N t u r i b i , (1979).
M e c h a n i c a l s t r e s s e s in f a s t l o c o m o t i o n of buffalo [Syncerus caffer) a n d e l e p h a n t
( L o x o d o n t a africana). Journal of Zoology 1 8 9 : 1 3 5 - 1 4 4 .
Biewener, A. A., J. T h o m a s o n , and L.E. L a n y o n , 1 9 8 3 . M e c h a n i c s of l o c o m o t i o n
a n d j u m p i n g in t h e f o r e l i m b of the h o r s e [Equus): In vivo s t r e s s d e v e l o p e d in t h e
r a d i u s a n d m e t a c a r p u s . Journal of Zoology 2 0 1 : 6 7 - 8 2 .
G r e e n b e r g , B. a n d / G . K. N o b l e , 1944. Social b e h a v i o r of t h e A m e r i c a n c h a m e l e o n
[Anolis carolinensis Voigt). Physiological Zoology 1 7 : 3 9 2 - 4 3 9 .
V
Dinosaur Necks and Tails
A LMOST ALL dinosaurs had long tails and some also had long necks.
In an extreme case, Diplodocus had a neck 7 meters (23 ft) long
(including the small head), a tail about 14 meters (46 ft) long and body
only 5 meters (16 ft) long. This chapter asks how dinosaurs used long
necks and tails.
I will start w i t h necks. T h e longest are those of sauropods, which
include the largest of all dinosaurs. Before the calculations about bone
strength had been done, m a n y people doubted w h e t h e r the biggest sau-
ropods could have supported their weight on land. They supposed that
they m u s t have lived in lakes, buoyed up by the water (figure 5.1).
Their long necks might have served as snorkels, enabling them to breathe
while standing on the b o t t o m in deep water.
The sauropods in figure 5.1 look like Diplodocus. If they are, their
lungs are 6 meters (20 ft) or more below the surface of the water. Their
necks are long enough to reach the surface but it would be very hard
for t h e m to breathe air in, because the lungs would have to be ex-
panded against a pressure difference of 6 meters of water. T h e snorkels
used by h u m a n divers are only about 30 centimeters (1 ft) long. The
dinosaurs in figure 5.1 would have needed e n o r m o u s chest muscles, to
breathe in.
It now seems clear that even the largest sauropods had legs strong
enough for walking on land, so there is no need to imagine t h e m living
submerged in lakes. Indeed, it has been argued that they probably lived
on dry land, keeping clear of marshy places where they could easily
have got bogged down because of the e n o r m o u s pressure on their feet
(chapter 3). If they lived on land, as is now generally believed, the long
neck cannot have been a snorkel. We m u s t look for some other func-
tion.
T h e shape of Brachiosaurus suggests that it lived like a giraffe, eat-
DINOSAUR NECKS A N D TAILS 6 1
FIGURE 5.1. Snorkcling sauropods, from Gregory 1951.
ing leaves from tall trees. N o t only was its neck very long, but the fore
legs were longer than the hind (a difference from other sauropods), so
its head could apparently be raised to the remarkable height of 13 me-
ters (43 ft). It was m u c h taller than any giraffe (figure 1.1).
If Brachiosaurus carried its head so high, its brain would be about 8
meters (26 ft) above its heart. T h e heart would have to p u m p blood at
very high pressure, to get it to the brain. If blood failed to get to the
brain even for a short time, the animal would collapse unconscious. A
fainting Brachiosaurus would come down w i t h a t r e m e n d o u s crash.
Blood pressure is conventionally measured in millimetres of mer-
cury because doctors used to use mercury m a n o m e t e r s to measure hu-
m a n blood pressure. It will be more convenient here to use meters of
water: a pressure of one meter of water equals 74 millimeters of mer-
cury. Modern reptiles p u m p blood out of their hearts at pressures that
are 0.5 to 1.0 meters of water above the pressure of the surrounding
tissues. This pressure difference is needed mainly to drive blood through
the capillaries, the fine blood vessels that permeate every tissue of the
body. Brachiosaurus would have needed an additional pressure of 8
meters of water to raise the blood to the brain, making a total of 8.5
or more meters of water. (Strictly speaking, the additional pressure would
be 8 meters of blood, not water, but this m a k e s little difference be-
cause the densities of blood and water are only slightly different.)
6 2 DINOSAUR NECKS AND TAILS
T h e calculated total blood pressure, 8.5 meters of water, is m u c h

larger than the blood pressure of any modern animal. Most m a m m a l s ,
including ourselves, p u m p blood from their hearts with pressures of
1.5 to 2.0 meters of water, and even giraffes do it with pressures no
higher than 4.3 meters of water. Giraffes need m u c h less blood pressure
than Brachiosaurus because their necks are so m u c h shorter, carrying
the brain only 3 meters above the heart.
You might think that the blood systems of giraffe and dinosaur necks
could work like siphons, which do not need high pressures to drive
liquids through t h e m . However, you need a rigid tube to make a siphon
because w h e n it is working, the pressure inside it, near the top, is less
than the pressure outside. Veins have flexible walls that would collapse
if the pressure inside fell, so they cannot work as siphons.
It would be a m i s t a k e to argue that a blood pressure of 8.5 meters
of water would be impossible, because it is so m u c h higher than the
blood pressures of modern animals. If giraffes were u n k n o w n , that kind
of argument would lead to the conclusion that giraffes were impossi-
ble. However, the blood pressure calculated for Brachiosaurus does seem
remarkable, and a very muscular heart would have been needed to pro-
duce it.
Brachiosaurus had long fore legs but Diplodocus and Apatosaurus
had relatively short ones, and look m u c h less like giraffes. In 1978 Dr.
Robert Bakker suggested that they also m a y have fed from tall trees,
rearing up on their hind legs to gain extra height (figure 5.2e,f). It may
seem ludicrous to suggest that these e n o r m o u s animals could manage
such gymnastics, but the idea deserves careful consideration. Remem-
F I G U R E 5.2. P o s s i b l e p o s t u r e s o f s o m e d i n o s a u r s w h e n b r o w s i n g o n f o l i a g e :
(a) Haplocanthosaurus; (b) Brachiosaurus ( n o t a v e r y l a r g e o n e ) ; (c) Camara-
saurus; (d)Barosaurus: (e) Diplodocus-, (f) Apatosaurus-, (g) Stegosaurus :a n d (h)
( f e e d i n g f r o m t h e g r o u n d ) Camptosaurus. F r o m B a k k e r 1 9 7 8 . R e p r i n t e d b y p e r -
m i s s i o n . C o p y r i g h t © 1978 M a c m i l l a n M a g a z i n e s Ltd.
DINOSAUR NECKS AND TAILS 63
ber that circus elephants can be trained to balance on their hind legs,
and that the calculations in chapter 4 suggest that Apatosaurus may
have been as athletic as elephants. T h e stresses in the bones, while
standing on the hind legs, would probably be less for Apatosaurus t h a n
for elephants. This is because in its normal quadrupedal position Apa-
tosaurus, unlike elephants, carried most of its weight on its hind legs
(table 4.1). Standing on the hind legs alone would not increase the load
on t h e m very m u c h .
It might be easy for Apatosaurus to support itself, once it was stand-
ing on its hind legs, but could it get itself into that position? It would
have to get its center of gravity over its hind feet, to take the load off
its fore feet. To discover w h e t h e r that would be difficult we need to
know where the center of gravity was. We already k n o w that Apato-
saurus and Diplodocus carried most of their weight on the hind legs,
which implies that the center of gravity was nearer the hind legs than
the fore. We could calculate its position from the data in table 4.1 but
we can discover it more directly, by a simple experiment.
For this I used the same models as in chapter 4, the solid plastic
ones with holes bored to represent the air-filled lungs. Their centers of
gravity should be in approximately the same positions as in the living
dinosaurs. I suspended each model by a thread tied to its head and
photographed it in side view (figure 5.3a). T h e model was hanging mo-
tionless, in equilibrium, so its center of gravity m u s t have been in line
with the thread, somewhere on the line AB. Then I suspended the model
from its back and took another photograph (figure 5.3b). T h e center of
gravity m u s t again have been in line w i t h the thread, on the line CD.
Finally, I superimposed the two photographs, m a k i n g the outlines of
the model coincide (figure 5.3c). T h e center of gravity was both on AB
and on CD: it m u s t have been at the intersection of the t w o lines.
Figure 5.4 shows centers of gravity located in this way. Diplodocus
standing in the position shown had its center of gravity over the left
hind foot. If the animal had moved its right hind foot forward and set
it down beside the left one, both hind feet would have been under the
center of gravity and it would have been easy for it to rear up on its
hind legs.
Diplodocus could apparently have reared up easily because its long,
heavy tail counterbalanced the front part of the body and brought the
center of gravity well back. Brachiosaurus had a shorter tail and heavier
fore quarters, so its center of gravity was further forward. It would have
been harder for it to rear up and, as far as I know, neither Bakker nor
anyone else has suggested that it did. With its long front legs and neck
it could feed from great heights w i t h o u t rearing up.
Bakker suggested that Stegosaurus reared up on its hind legs to feed,
64 DINOSAUR NECKS A N D TAILS
and it seems likely that it could have done: its center of gravity was
well back, near the hind legs. Triceratops had its center of gravity much
further forward (figure 5.4) and might have found it difficult to rear up.
However, elephants have their centers of gravity well forward, and can
rear up. (Table 4.1 shows that the centers of gravity of elephants are
far enough forward to put most of the weight on the front feet in nor-
mal standing.)
It is generally assumed that long-tailed sauropods such as Diplodo-
cus and Apatosaurus walked around with their necks nearly horizon-
tal. Most drawings show t h e m in that position, as do the m o u n t e d
skeletons in m u s e u m s . We will think about the problem of supporting
a long, heavy, horizontal neck.
I am going to suggest that these necks were supported in the same
way as the necks of horses, cattle, and their relatives. These animals
have a thick ligament called the l i g a m e n t u m nuchae running along the
backs of their necks (figure 5.5). Unlike most other ligaments it con-
sists mainly of the protein elastin, which has properties very like rub-
ber. It can be stretched to double its initial length without breaking
and snaps back to its initial length w h e n released.
The ligamentum nuchae is stretched when the animal lowers its head
F I G U R E 5.3. D i a g r a m s s h o w i n g h o w to l o c a t e t h e c e n t e r of gravity of a m o d e l .
T h e y are explained in the text.
DINOSAUR NECKS AND TAILS 65
F I G U R E 5.4. O u t l i n e s of dinosaurs, s h o w i n g t h e positions of their c e n t e r s of

gravity.
to drink or graze, and shortens again w h e n the head is raised. In ex-

periments with deer carcasses, my colleagues and I found that the lig-
ament was 1.4 times its slack length-when the head was raised to the
position of figure 5.5a, and almost twice its slack length w h e n it was
lowered to the position of figure 5.5b. Notice that the ligament was
stretched even w h e n the head was high: I doubt w h e t h e r a deer can
get into a position that allows the ligament to shorten to the point of
going slack. If you cut the ligament in a dissection the cut ends spring
apart, as if you had cut a stretched rubber band.
T h u s the l i g a m e n t u m n u c h a e is taut in all normal positions of the
neck. Its tension helps to support the weight of the head and neck.
The tension and the supporting effect are greatest w h e n the head is
lowered (as in figure 5.5b) but even in this position the tension is not
by itself enough to give all the necessary support: some tension in neck
muscles is needed as well.
The ligamentum nuchae is a feature of hoofed m a m m a l s but some-
thing rather similar is found in birds, w h i c h are m u c h more closely
related to dinosaurs. Instead of a continuous ligament running the whole
length of the neck they have a series of short ligaments connecting
each neck vertebra to the next (figure 5.6a). These ligaments consist
largely of elastin, like the ligamentum nuchae. They are stretched when
the head is lowered, and shorten again w h e n it is raised.
Figure 5.6c shows a bird vertebra in front view. T h e c e n t r u m is the
main body of the vertebra, connected to the vertebrae in front and be-
hind by intervertebral discs. Above it is a hole for the nerve cord and
above that again is the neural spine. T h e elastic ligaments connect the
front of one neural spine to the back of the next.
66 DINOSAUR NECKS AND TAILS
Diplodocus and Apatosaurus have neck vertebrae w i t h V-shaped

neural spines (figure 5.6d). I suggest that the V was filled by an elastin
ligament that ran the whole length of the neck and back into the trunk.
This ligament would have helped to support the neck while allowing
the dinosaur to raise and lower its head.
I made some experiments to find out w h e t h e r the idea was feasible.
I cut off the head and neck of the Diplodocus model and measured
their volume (as I had done w i t h complete models in chapter 2), and
calculated that the mass of the real head and neck would have been
1,340 kilograms. Their weight was this mass multiplied by the accel-
eration of gravity, 1,340 x 10 = 13,400 newtons. I suspended the am-
putated head and neck from threads to find their center of gravity, and
have shown the weight acting at the center of gravity in figure 5.6b.
We can only guess how thick the ligament was, but it seems likely
that it projected above the tops of the neural spines, as shown in figure
5.6d. If so its center line, at the base of the neck, was about 0.42 meter
above the center of the c e n t r u m .
In figure 5.6b the weight is pulling the neck counterclockwise about
the joint at its base and the tension in the ligament is pulling it clock-
wise. T h e weight acts 2.2 meters from the joint and the ligament ten-
sion 0.42 meters from it. By the principle of levers, the tension that
would be needed to balance the weight is 2.2 x 13,400/0.42 = 70,000
F I G U R E 5.5. T h e n e c k o f a R o e d e e r i n t h e a l e r t p o s i t i o n a n d l o w e r e d for feed-

ing. T h e s k e l e t o n a n d t h e l i g a m e n t u m n u c h a e (stippled) are s h o w n .
DINOSAUR NECKS AND TAILS 6 7
F I G U R E 5.6. N e c k s o f (a) a t u r k e y a n d (b) Diplodocus, s h o w i n g t h e v e r t e b r a e

a n d t h e e l a s t i c l i g a m e n t s . T h e f o r c e s t h a t a c t e d o n t h e n e c k o f Diplodocus are
a l s o s h o w n , (c) a n d (d) a r e f r o n t v i e w s o f n e c k v e r t e b r a e o f a n o s t r i c h a n d o f
Diplodocus.
newtons (7 tonnes force). T h e third force s h o w n in the diagram is the

force in the joint itself, where one c e n t r u m presses on the next.
The calculated tension may seen enormous, but the ligament was
very thick. If it was as thick as in the diagram, its cross-sectional area
was 40,000 square millimeters and the stress in it, for a force of 70,000
newtons, would be 1.8 n e w t o n s per square millimeter. This is more
than the stress in the l i g a m e n t u m n u c h a e of a deer w i t h its head down
(about 0.6 newtons per square millimeter), and would be enough, or
nearly enough, to break ligamentum nuchae. However, this stress would
act only if the ligament supported the neck w i t h o u t any help from
muscles. If neck muscles took some of the load (as they do in birds)
the stress in the ligament would be less. T h e suggestion of an elastin
ligament seems feasible.
Remember that the ligament is only a guess, based on the structure
of sauropod vertebrae and comparisons w i t h modern animals. We do
not know whether it existed, but the calculations seem to show that
if it did it could have done a useful job. Only some sauropods had V-
shaped neural spines that could have housed a continuous ligament but
other dinosaurs may have had separate ligaments connecting each neck
vertebra to the next, as in birds.
N o w we will think about tails. We have already seen how a long tail
may balance the front part of the body, enabling some dinosaurs to rear
68 DINOSAUR NECKS AND TAILS
up on their hind legs. T h a t function depends on the animal being able

to lift its tail off the ground. If the tail lay limp on the ground, m u c h
of its weight would be supported directly on the ground and it would
be little use as a counterpoise. M a n y drawings of dinosaurs show the
tail trailing on the ground but few sets of footprints show the mark
that would be made by a trailing tail (chapter 3). It seems possible that
the thin end of the very long tail of Diplodocus trailed on the ground,
but this would have little effect on the counterpoise function if this
part of the tail was as thin as the slender vertebrae suggest.
Though Diplodocus would have to be able to stiffen its tail to raise
it for use as a counterpoise, it would also have to be able to bend its
tail to get into the position s h o w n in figure 5.2. T h e tails of ornitho-
pods may have been m u c h stiffer. They have a crisscross arrangement
of rods, on either side of the neural spines of their vertebrae, that seem
to be ligaments or tendons turned to bone (figure 5.7). (You will find
tendons that have turned to bony material whenever you eat turkey,
as hard strips embedded in the meat of the lower leg.) Whenever com-
plete skeletons of ornithopods are found, the tail is fairly straight (fig-
ure 5.7), suggesting that it was indeed stiff.
How stiff the tail was would depend on w h e t h e r the rods were lig-
a m e n t s or tendons. Ligaments connect bone to bone, and if the tail
vertebrae were connected by rigid bony ligaments the tail would be
very stiff indeed. Tendons connect muscles to bones, and if the rods
were tendons the tail could have bent up and down a little as its m u s -
cles shortened and lengthened. T h e rods look like tendons to me. Mod-
ern m a m m a l s have similarly arranged (but non-bony) tendons in their
tail muscles.
Bony ligaments or tendons are m o s t p r o m i n e n t in the duck-billed
F I G U R E 5 . 7 . S k e l e t o n o f Iguanodon, i n t h e p o s i t i o n i n w h i c h i t w a s f o u n d .
N o t i c e t h e crisscross t e n d o n s i n t h e b a c k and tail. F r o m N o r m a n 1980.
DINOSAUR NECKS A N D TAILS 69
F I G U R E 5 . 8 . A n o s t r i c h , a m a n , a n d a s m a l l b i p e d a l d i n o s a u r [Hypsilophodon]
running, s h o w i n g the positions of their hips and their centers of gravity: O h i p ;
• c e n t e r of g r a v i t y .
dinosaurs but have also been found over the hips of Triceratops, in the
tail of the theropod Deinonychus and in several other dinosaurs.
The tail m u s t have had a major effect on the running m o v e m e n t s of
bipedal dinosaurs. Figure 5 . 8 compares a dinosaur w i t h two modern
bipeds, a bird and a h u m a n . T h e dinosaur, w i t h its long heavy tail, has
the center of gravity close to the hips, but the bird, w i t h only a tuft
of feathers for a tail, has its center of gravity well in front of the hips.
The h u m a n has no tail but the center of gravity is close to the hips
because the trunk is erect. If these bipeds are not to fall over, the av-
erage positions of their feet, while on the ground, m u s t in each case
be under the center of gravity. Each animal sets the foot down in front
of the center of gravity and does not lift it u n t i l it is behind the center
of gravity. The bird manages by holding its thigh almost horizontal and
swinging the leg from the knee. It seems likely that bipedal dinosaurs
moved their legs more like people than like birds, because of the po-
sition of the center of gravity.
The tails of kangaroos are thick and heavy, but they are flexible, and
they swing up and down as the animal hops. If flexible tails are suitable
for kangaroos, why should stiff ones have evolved in bipedal dinosaurs?
The answer may depend on the difference between running and hop-
ping and on a basic principle of mechanics.
I will use an example from gymnastics to explain the principle of
conservation of angular m o m e n t u m . A gymnast on a trampoline can
set his body spinning, while in mid air, by moving his arms. By swing-
ing his arms to the left he m a k e s his body spin to the right. A rotating
object has a property called angular m o m e n t u m that depends on the
masses of its parts and on its rate of rotation. T h e principle says that
the gymnast cannot change his total angular m o m e n t u m without

pushing on something, but he can set his arms rotating in one direction
and his t r u n k in the other so that the angular m o m e n t u m s in the two
directions cancel out.
As a kangaroo flies through the air in a hop, it swings its legs forward
ready for the next landing. In figure 5.9 its legs have to be given coun-
terclockwise angular m o m e n t u m so its trunk and tail must get match-
ing clockwise m o m e n t u m . If the tail were rigid the body would rock
up and down quite a lot (through 13-18°, according to my calcula-
tions]. Actually the tail bends so that it swings up and down through
a large angle and the trunk through a m u c h smaller one. These tail
m o v e m e n t s probably cost the animal very little energy, because they
do not have to be powered by muscles. T h e tail vibrates passively be-
cause its long fnon-bony) tendons m a k e it springy.
If the trunk rocked a lot, with the head rocking with it, it might be
difficult for the animal to keep watch for danger as it hopped. T h e
springy tail m a y give the kangaroo an advantage, by reducing the rock-
ing m o v e m e n t s of the trunk. A dinosaur that ran rather than hopped
would not need such a m e c h a n i s m because, in running, one leg swings
back while the other swings forward and there is little tendency for
leg m o v e m e n t s to rock the body. A springy tail m a y be best for a hop-
per but a stiff one seems fine for a runner.
T h e tails of some dinosaurs may have served as weapons, as well as
for balance. It has often been suggested that the long tapering tails of
Diplodocus and Apatosaurus would have made formidable whips. They
could have been used to strike a predator, and I wonder whether they
could also have been used to m a k e a terrifying noise. When a circus
ringmaster cracks his whip, he flicks it so as to m a k e its tip move
supersonically. Is it too wild a speculation to wonder whether Diplo-
docus could crack its tail?
F I G U R E 5.9. D i a g r a m of a k a n g a r o o h o p p i n g .
DINOSAUR NECKS AND TAILS 7 1
(b)
FIGURE 5.10. Euoplocephalus o r Dyoplosaurus, a n a n k y l o s a u r w i t h a c l u b o n

i t s t a i l (a) i n s i d e v i e w a n d (b) i n t o p v i e w . L e n g t h s i x m e t e r s . F r o m C a r p e n t e r
(1982).
There are other dinosaur tails that seem more obviously to have been
weapons. Stegosaurus had sharp spikes on its tail up to half a meter
(20 in) long. Some of the ankylosaurs had big l u m p s of bone at the ends
of their tails and presumably used t h e m as clubs (figure 5.10).
This chapter has told a series of short stories. Sauropods are unlikely
to have used their long necks as snorkels. Very large breathing muscles
would be needed for snorkeling at any substantial depth, and in any
case it seems unlikely that they lived in water. It seems more likely
that they lived on land, raising their necks to feed from high branches.
Their hearts would then have had to p u m p blood out at high pressure,
to get it to the brain. Brachiosaurus probably fed like a giraffe but Di-
plodocus and similar sauropods may have reared up on their hind legs
to get their heads high. Their long, heavy tails would probably have
made it easy for these sauropods to get their hind feet under their cen-
ter of gravity. The long necks of Diplodocus and Apatosaurus may have
been supported by an elastin ligament running through the V-shaped
notches in their neural spines.
The tails of duck-billed dinosaurs had bony tendons or ligaments
alongside the neural spines and seem to have been stiff. T h e effect of
the tail on the position of the center of gravity probably made bipedal
dinosaurs move their legs more like people than like birds. T h e springy
tails of kangaroos may help to prevent the body from rocking too m u c h
during hopping but stiff tails seem suitable for running bipeds. The
tails of some dinosaurs seem to have served as weapons.
Principal Sources
C h a p t e r 5 , l i k e c h a p t e r 4 , i s b a s e d largely o n A l e x a n d e r (1985), w i t h s o m e o f t h e
e x p e r i m e n t s m o d i f i e d . T h e d a t a a b o u t blood p r e s s u r e are from H o h n k e (1973). T h e
s u g g e s t i o n t h a t s o m e s a u r o p o d s m a y h a v e reared u p o n t h e i r h i n d legs w a s m a d e
by B a k k e r (1978). T h e e l a s t i n l i g a m e n t s in t h e n e c k s of birds w e r e i n v e s t i g a t e d by
B e n n e t t a n d A l e x a n d e r (1987). G a l t o n (1970) d e s c r i b e d t h e stiff tails of d u c k - b i l l e d
d i n o s a u r s . A l e x a n d e r a n d V e r n o n (1975) d i s c u s s e d t h e o s c i l l a t i o n s of t h e t a i l s of
kangaroos.

Zoological journal of the Linnean Society 8 3 : 1 - 2 5 .
A l e x a n d e r , R. M c N . a n d A. V e r n o n , 1975. T h e m e c h a n i c s of h o p p i n g by k a n g a r o o s
( M a c r o p o d i d a e ) . journal of Zoology 1 7 7 : 2 6 5 - 3 0 3 .
Bakker, R. L. 1978. D i n o s a u r feeding b e h a v i o r and t h e origin of f l o w e r i n g p l a n t s .
Nature 2 7 4 : 6 6 1 - 6 6 3 .
B e n n e t t , M. B. a n d R. M c N . A l e x a n d e r , 1987. P r o p e r t i e s a n d f u n c t i o n of e x t e n s i b l e
l i g a m e n t s in t h e n e c k s of t u r k e y s [Meleagris gallopavo) a n d o t h e r birds, journal
of Zoology 2 1 2 : 2 7 5 - 2 8 1 .
C a r p e n t e r , D. 1982. S k e l e t a l a n d d e r m a l a r m o r r e c o n s t r u c t i o n of Euoplocephalus
lulus. Canadian journal of Earth Sciences 1 9 : 6 8 9 - 6 9 7 .
G a l t o n , P. M. 1970. T h e p o s t u r e of h a d r o s a u r i a n d i n o s a u r s , journal of Paleontology
44:464- 473.
G r e g o r y , W. K. 1 9 5 1 . Evolution Emerging. N e w York: M a c m i l l a n .
H o h n k e , L. A. 1973. H a e m o d y n a m i c s in t h e s a u r o p o d s . Nature 2 4 4 : 3 0 9 - 3 1 0 .
N o r m a n , D. B. 1980. On t h e o r n i t h i s c h i a n d i n o s a u r Iguanodon bernissartensis of
B e r n i s s a r t (Belgium). Memoires de l'Institut Royal des Sciences Naturelles de Bel-
gique 1 7 8 : 1 - 1 0 3 .
VI
Fighting and Singing Dinosaurs
r n H E T A I L S mentioned at the end of chapter 5 are not the only weap-
X ons that dinosaurs had. I will n o w tell about horns and reinforced
heads, and about how they m a y have been used for fighting. I also spec-
ulate about how dinosaurs m a y have approached rivals, and about the
noises they may have made.
It seems likely that male dinosaurs fought rival males of the same
species, m u c h as stags fight w i t h their antlers and male antelope fight
with their horns. It may seem odd that animals fight, getting h u r t and
possibly killed. You may t h i n k that peaceful species should flourish
and quarrelsome ones should destroy themselves, and that evolution
by natural selection should m a k e all species peaceful.
To understand why it does not, you need to understand the principle
of the survival of the fittest. This principle is not about the survival
of species or even the survival of individual animals: it is about the
survival of genes, the basic u n i t s of heredity. Genes are complex mol-
ecules that carry information. Some carry patterns for m a k i n g other
molecules. Other genes or groups of genes carry instructions for mak-
ing parts of the body, or for patterns of behavior. Almost every animal
(there are some exceptions) inherits half its genes from its m o t h e r and
half from its father. In turn, it passes on copies of just half of its genes
in each egg or spermatozoon that it m a k e s . T h u s genes are transmitted
from generation to generation. Errors of copying produce new genes
from time to time, and the mixing of genes that happens in reproduc-
tion brings new combinations of genes together. Some genes or groups
of genes m a k e their possessors more successful in reproduction, and
are more likely than others to be passed on to successive generations.
Some of t h e m are obviously beneficial. For example, a gene that made
its possessors i m m u n e to disease, or enabled t h e m to run faster to es-
cape predators, would m a k e t h e m better able to survive and reproduce.
74 FIGHTING AND SINGING DINOSAURS
It would be more likely than alternative genes, that did not give these
advantages, to be passed on to successive generations. Once such a gene
had appeared it would be likely to become c o m m o n e r until, m a n y gen-
erations later, all m e m b e r s of the population had it.
O t h e r genes that are less obviously beneficial will also increase. Sup-
pose that a gene made its male possessors very aggressive, quick to grab
and copulate with every female they m e t and to chase other males
away. Males with that gene would be apt to get involved in a lot of
fights, to get hurt and to die young, but they might also get unusually
m a n y offspring. If they did, the gene would be favored by natural se-
lection.
It is m u c h more likely that males will fight for females than that
females will fight for males. T h e reason is that females have to put a
lot of energy and materials into m a k i n g each egg or embryo, so they
can produce only a limited n u m b e r of offspring. Males, however, can
produce e n o r m o u s n u m b e r s of sperm. A female can get all her eggs
fertilized by a single father but a male has plenty of sperm for many
mates. If the n u m b e r s of males and females are about equal (as they
are for most species) the females will easily get all the matings they
can use but the males will have plenty of sperm to spare. The males
could get more offspring (and pass more genes on to future generations)
if they could copulate with more females and keep other males away
from t h e m .
Triceratops has horns that m u s t surely have been weapons (figure
1.13). It m u s t have looked rather like a huge rhinoceros with a peculiar
arrangement of horns, but these are more like antelope than rhinoceros
horns. Antelopes have spikes of bone covered w i t h horn but rhinoceros
horns are consolidated tufts of hair with no bony core. Triceratops horns
are spikes of bone and were presumably covered w i t h horn when the
dinosaur was alive.
Many of the large, plant-eating m a m m a l s have horns of some kind.
Antelopes, cattle, and sheep have true, p e r m a n e n t horns, and deer have
antlers which they shed and re-grow annually. All these animals may
s o m e t i m e s use their horns to defend themselves against predators, but
their usual defense is to run away. T h e principal use of their horns and
antlers seems to be for fighting between males of the same species.
Red deer are a European species, closely related to the American
wapiti. Their stags assemble harems, usually of about six females but
s o m e t i m e s of twenty, and defend t h e m from other males. Only stags
that have harems breed, but each of t h e m may father many offspring.
Genes that help stags to get and keep large harems m u s t be favored by
natural selection.
Stags that have no harem try to get one and those that have a harem
FIGHTING AND SINGING DINOSAURS 7S
already try to add to it. T h i s leads to fighting between rival males.

They run at each other and clash antlers. T h e antlers interlock (figure
6.1a) and the stags wrestle, each trying to push the other back or to
throw it to the ground. A stag that manages to throw another m a y stab
it with the points of its antlers, but the loser usually gives up and runs
away without being thrown. Antelopes interlock horns and wrestle in
the same way.
Many other species of deer defend harems, as do m a n y antelopes,
but other animals with different social systems fight in different cir-
cumstances. Bighorn sheep do not keep harems, but males seek out
females that are in estrus and guard just one female at a t i m e . They
stay w i t h her for one to three days, while she remains in estrus, cop-
ulating about once an hour to m a k e sure that their sperm are in the
right place at the right time, w h e n the female ovulates. O t h e r males
try to capture estrus females from males that are guarding them, or to
sneak a quick copulation while the guarding male is distracted. This
leads to fighting. Sheep horns do not interlock, so there is no wrestling.
Bighorns and other wild sheep fight by running at each other and col-
liding head-on, or by rearing up and clashing horns (figure 6.2).
If two Triceratops ran at each other w i t h their heads down, their
horns would interlock (figure 6.1b). It seems likely that males inter-
F I G U R E 6 . 1 . (a) R e d d e e r s t a g s w i t h t h e i r a n t l e r s i n t e r l o c k e d , d r a w n f r o m a
p h o t o g r a p h i n C l u t t o n - B r o c k 1 9 8 2 ; (b) Triceratops f i g h t i n g w i t h t h e i r h o r n s i n -
terlocked, d r a w n from photographs of m o d e l s sold by the N a t u r a l History M u -
seum, London.
FIGURE 6.2. (a) T h i n h o r n r a m s f i g h t i n g , f r o m G e i s t 1 9 7 1 ; (b) Stegoceras fight-

ing.
locked horns and wrestled like stags and antelopes, but it seems just
a little doubtful w h e t h e r their horns were strong enough.
Figure 6.3 shows that Triceratops had rather small horns for its size,
by antelope standards. Horn reach (figure 6.3a) m e a n s the straight-line
distance from horn base to horn tip. T h e reach of a 6-tonne Triceratops
is little different from that of an eland of one-tenth its mass, and far
less t h a n w h a t the lines suggests 6-tonne antelope would have, if an-
telope grew so big. However, the shortness of the horns might not mat-
ter, provided they were strong enough.
Figure 6.3b shows the cross-sectional areas of the bony cores of horns,
at the horn base. Female antelope have thinner horns than males of
the same species, or no horns at all (but w h e n they have horns they
are generally about as long as those of males). T h e graph shows that
Triceratops horns are t h i n n e r than would be expected on antelopes of
the same mass, w h e t h e r male or female. T h a t m e a n s that Triceratops
horns were rather weak, for the size of animal.
We need to be rather careful about the argument here, because short
horns may not need to be as strong as long ones would have to be on
the same animal. R e m e m b e r the passage in chapter 4 where I said that
long thin structures such as leg bones or horns are most easily broken
by forces acting at right angles to t h e m , which set up bending mo-
m e n t s in t h e m . T h e m a x i m u m stress at a distance x from a force R
acting at right angles to the horn is Rx/Z, where Z is the quantity
called the section m o d u l u s . A thin horn has a low section modulus
FIGHTING AND SINGING DINOSAURS 77
but, if it is also short, it has low x as well as low Z. T h e stress set up

in it by a force R may be no more than in a thicker, longer horn.
We might think that the shortness of Triceratops horns made up for
their thinness, but for two things. First, wrestling antelopes engage the
bases of their horns, not the tips, so x m a y not be proportional to horn
length. Second, figure 6.3 shows little sign of a trade-off between thick-
ness and length. The oryx has remarkably long horns that are also rather
thin for the size of antelope and the wildebeest has short horns that
are rather thick. There is no question of horns being so strong that
differences of thickness do not matter: 3 percent of male wildebeest
and 17 percent of male oryx, counted in wild populations in East Af-
rica, had one or both horns broken. Triceratops horns seem rather weak,
for so large an animal.
You might suppose that horns used mainly for fighting between males
would be grown only by males. Antlers are indeed grown only by male
deer, with one exception: both sexes of caribou have them. Horns,
however, are grown by both sexes of m a n y antelope species, as figure
6.3 shows, and also by both sexes of cattle and sheep. If ceratopians
used their horns mainly for fighting between males you should not
necessarily expect to find horns in males only, but you might reason-
ably expect the horns to be thicker in males than in females. T h e fos-
sils show no clear signs of this. So far, scientists have been unable to
decide which are the male skulls and which the female ones, for any
of the horned ceratopian species. (They think they have worked it out
for Protoceratops, which has no horns.)
When a male deer or antelope challenges another, they do not fight
immediately. Each puts on a display which enables the other to gauge
its strength. In the case of Red deer the first stage is a roaring m a t c h .
Each stag roars loudly and repeatedly. As the contest continues, roars
follow each other at shorter and shorter intervals. If the challenging
stag finds he cannot manage as m a n y roars per m i n u t e as his opponent,
he may give up and go away: roaring rate seems to be used as an in-
dicator of strength. If he does not give up at this stage the stags walk
side by side for a while, looking at each other and judging each other's
strength. Again, the challenger may give up, but otherwise the stags
fight.
The elaborate ritual before fights seems to have evolved for a good
reason. A belligerent stag whose genes made h i m fight for females on
all possible occasions might be tremendously successful in reproduc-
tion if he happened to be strong enough to win every fight. More prob-
ably, he would often lose fights. He might get badly beaten up while
still quite young and leave very few offspring.
N o w imagine a different set of genes that m a k e s stags behave in a

m u c h more calculating way, that m a k e s t h e m fight only when they
think they can win and run away when they are sure they would lose.
Stags with these genes are likely to live longer and leave more offspring
than stags that fight indiscriminately. T h e better they are at judging
strength the more offspring they are likely to leave, but even if their
judgment is not very good it should help. T h e set of genes is likely to
be favored by natural selection. T h a t is w h y Red deer have evolved the
strength-assessing ritual. It seems likely that ceratopians did some-
thing rather similar.
Figure 6.4 shows the frill at the back of Triceratops' head, as well as
the horns. This frill was formed by a strong sheet of bone, extending
T h e a n t e l o p e d a t a a r e f r o m P a c k e r 1 9 8 3 . T h e Triceratops h o r n m e a s u r e m e n t s
a r e f r o m H a t c h e r 1 9 0 7 , a n d t h e Triceratops m a s s i s f r o m c h a p t e r 2 .
backward from the skull. There seem to have been big jaw muscles
whose a t t a c h m e n t s extended onto the bony frill, but the frill seems
too large to have evolved just for that. It m u s t have been a useful shield,
protecting the animal's neck from rivals' horns, but some other cera-
topian frills were supported by bone only round the edge and would
have been less good as shields unless their skin was very thick. T h e
frill of Styracosaurus w i t h its splendid row of spikes seems designed
as much for show as for protection (figure 6.4).
Perhaps we can find a parallel among deer. Red deer have fairly prac-
tical-looking antlers: the spikes are weapons and the branches (espe-
cially the ones over the eyes) give protection by catching rivals' antlers.
Moose, Fallow deer, and some others have palmated antlers, incorpo-
so FIGHTING AND SINGING DINOSAURS
F I G U R E 6 . 4 . S k u l l s o f Triceratops a n d Styracosaurus, drawn from illustra-

t i o n s in H a t c h e r 1907 a n d B r o w n and Schlaikjer 1937.
rating broad plates of bone that look most impressive but seem un-
necessary for any mechanical function. They may help to attract fe-
males, like the long tails of male widowbirds which I discuss later in
this chapter. Female Red deer m a t e w i t h the stags that win t h e m but
Moose and Fallow deer have different social systems, and their females
may have more choice of mate. Whatever the function of the bony
plates on palmated antlers, the function of the frill of Styracosaurus
was probably similar.
N o w we will look at another probable weapon, the thick skull roof
of the pachycephalosaurs (also called dome-headed dinosaurs). Figure
6.5 shows the skull of one of these animals [Stegoceras] and, for com-
parison, the skull of a similar-sized ornithopod. T h e huge bulge on the
top of Stegoceras' skull is solid bone. Though the skull is only 28 cen-
timeters long its roof is eight centimeters thick. Stegoceras is a small
dinosaur, about two meters long, but some other pachycephalosaurs
were m u c h larger.
Their thick skull roofs look like battering rams. It seems likely that
male pachycephalosaurs fought like Bighorn sheep, running at each other
with heads down (figure 6.2). The thick skull roofs would take the im-
pact, like the horns of the sheep. Some pachycephalosaur skulls have
thinner roofs than others that seem to be the same species, and may
be female. Similarly among sheep, ewes have smaller horns than rams.
We will think about the forces that might have acted when pachy-
cephalosaurs collided. To simplify the calculation, imagine t w o iden-

tical dinosaurs running at equal speeds, colliding head-on. If everything
is equal, neither is pushed back by the other: it is as if each collided
with an absolutely rigid wall. If the dinosaur itself were rigid, it would
be stopped instantly, but that would need an infinite force, which is
impossible. The dinosaur would have to deform a little.
A running dinosaur has kinetic energy, which it loses w h e n it stops.
This energy equals half its mass t i m e s the square of its speed
kinetic energy = fa mass x (speed)

l 2
The energy absorbed in the impact is the force multiplied by the de-
celeration distance
energy absorbed = force x deceleration distance.
To stop the dinosaur, all the kinetic energy m u s t be absorbed, so

L
/ 2 mass x (speed) = force x deceleration distance
2
mass x (speed) 2
force =
2 x deceleration distance
If the mass is measured in kilograms, the speed in meters per second
and the distance in meters, this equation gives the force in newtons.
The less rigid the dinosaur, the more it deforms, the bigger the deceler-
F I G U R E 6.5. S k u l l s of Stegoceras (a p a c h y c e p h a l o s a u r ] a n d Hypsilophodon ( a n

ornithopod), from Sues 1978.
ation distance and so (from the equation) the smaller the force. The
same is true of colliding cars, which is why crumple zones save lives.
(Crumple zones are parts of a car body designed to crumple in an impact.)
A crumple zone is good for just one crash, but pachycephalosaurs
presumably clashed heads often. What they would have needed, to
moderate the forces, is some sort of elastic padding that would deform
in the impact but spring back into shape afterward. The bulging skull
roof m a y have been quite effective as padding. Sections cut through it
show that the bone was slightly spongy, suggesting that it could have
deformed quite a lot in an impact.
I want you to realize just how big the forces might have been, if the
dinosaur were too rigid. Imagine a 20-kilogram dinosaur (a very rough
estimate of the mass of Stegoceras) running at 3 meters per second.
This would be a slow jogging speed for a h u m a n , and is well within
the range of speeds estimated from dinosaur footprints in table 3.2.
Suppose that dinosaur were in a collision that stopped it in a distance
of one centimeter or 0.01 m e t e r (it is hard to imagine the skull roof
deforming more). T h e force would be (20 x 3 )/(2 x 0.01) = 9,000 new-
2
tons, or 0.9 tonnes force. The skull roof of Stegoceras looks strong enough
for that but I doubt w h e t h e r the neck vertebrae could have stood it.
(Unfortunately, no neck vertebrae have been found and we can only
guess their strength from the sizes of trunk vertebrae.)
The double lines in figure 6.5 show the angles at which the neck
vertebrae seem to have joined the skull. (This can be judged from the
shape of the a t t a c h m e n t area for the first vertebra, on the back of the
skull.) T h e angle between the skull and the neck of Stegoceras means
that, w h e n the head was down in the butting position, the neck would
have been more or less in line w i t h the force. T h e trunk vertebrae of
pachycephalosaurs interlock in a way that seems likely to have made
the spine rather rigid, and there were bony tendons or ligaments in the
back as well as the tail. Scientists have interpreted these signs as show-
ing that pachycephalosaurs had very stiff backs and have pictured t h e m
colliding w i t h their backs ramrod straight. I find that hard to believe
because the forces (for any likely r u n n i n g speed) would be so large.
They would be m u c h smaller if the backbone buckled at impact; the
skull would still be decelerated over a very short distance but most of
the mass of the body would travel further forward, before being stopped.
T h e neck could serve as a crumple zone, but one that could be straight-
ened after the impact and used again.
Some of the hadrosaurs (duck-billed dinosaurs) also have strange pro-
jections from their skulls. Anatosaurus (figure 1.11) has a relatively
plain skull but some other hadrosaurs with similar-shaped bodies have
extraordinary crests (figure 6.6). Corythosaurus' crest is an upward-
F I G U R E 6 . 6 . S k u l l s of c r e s t e d h a d r o s a u r s (a] Corythosaurus; (b) Lambeosau-

rus a n d (c| Parasaurolophus. T h e s c a l e l i n e s a r e 2 0 c m l o n g . F r o m W e i s h a m p c l
:
1981.
pointing semicircle, Lambeosaurus' crest has two branches and Para-

saurolophus' crest is a backward-pointing rod. Other hadrosaur crests
had other shapes. They do not look like weapons and they cannot have
been very strong: some hadrosaurs had small solid crests but all the
ones in figure 6.6 are hollow, with extensions of the nose cavity inside.
Why did they evolve?
It seems likely that they evolved for the same reason as cocks' combs
and peacocks' tails, as conspicuous o r n a m e n t s . A m o n g hadrosaurs that
seem to belong to the same species there are some with large crests
that were probably male and others w i t h smaller crests that were prob-
ably female. Similarly, cocks have bigger combs (and tails) than hens,
and peacocks have i m m e n s e l y longer, h a n d s o m e r tails t h a n peahens.
These structures seem to have evolved by a process called sexual
selection: they evolved because females preferred highly ornamented
males. T h e best evidence that this can happen comes from experiments
with Long-tailed widowbirds in Kenya. These birds are about the size
of American robins. The females are brown and inconspicuous but the
males are black w i t h extremely long (half meter) tails. They live in
grassland, where the males establish territories averaging about a hec-
tare (the area of a baseball outfield) and defend t h e m against other males.
They advertize themselves to females by flying low over their terri-
tories with their tails spread. Each female apparently selects a male,
mates with him, and nests on his territory. Like Red deer stags, suc-
cessful males m a t e w i t h several females, but they do not compete for
females by fighting each other. Instead, they compete to attract fe-
males.
Malte Andersson, a Swedish zoologist, did an experiment to test the
hypothesis that longer tails are more attractive to females. He caught
male widowbirds, altered the lengths of some of their tails, and re-
leased t h e m again. He cut some of their tails down to one third of their
8 4 FIGHTING AND SINGING DINOSAURS
original length and used the cut-off feathers to lengthen the tails of
other males, by sticking t h e m on w i t h superglue. He left other birds
with normal-length tails, either leaving t h e m intact or cutting t h e m
off and sticking t h e m back on again to check for possible effects of
cutting and gluing. Before the experiment, each group had an average
of 1.5 nests per territory (figure 6.7a), but m a n y of the females had not
yet mated. During the following m o n t h , males w i t h elongated tails
gained an average of 1.8 additional nests each, but the other groups of
males gained only 0.5-1.0 additional nests per territory (figure 6.7b).
T h e unnaturally elongated tails seem to have attracted females. This
F I G U R E 6 . 7 . W i d o w b i r d s w i t h l o n g t a i l s a t t r a c t m o s t f e m a l e s : (a) t h e m e a n
n u m b e r s o f n e s t s p e r t e r r i t o r y for f o u r g r o u p s o f m a l e s , b e f o r e t h e e x p e r i m e n t ;
(b) t h e m e a n n u m b e r s o f e x t r a n e s t s b u i l t i n t h e m o n t h a f t e r t h e i r t a i l s h a d
b e e n s h o r t e n e d , e l o n g a t e d , c u t a n d g l u e d b a c k o n ( c o n t r o l I ) o r left u n a l t e r e d
( c o n t r o l II). F r o m A n d e r s s o n 1 9 8 2 . R e p r i n t e d b y p e r m i s s i o n . C o p y r i g h t © 1 9 8 2
M a c m i l l a n M a g a z i n e s Ltd.
FIGHTING AND SINGING DINOSAURS
suggests that male widowbird tails evolved to their normal (already

remarkable) length because the females liked t h e m long. T h e normal
length is probably a compromise between female preference and the
inconvenience of an excessively long tail. Similarly, hadrosaurs prob-
ably evolved their crests because females liked t h e m .
That leaves the question, why should females prefer anything so bi-
zarre? Suppose you were a female hadrosaur w h o did not care for big
crests, while most other females thought t h e m marvelously sexy and
desirable. You may think that a preference for big crests is just an ir-
rational fad, but if you breed w i t h a small-crested male your sons will
probably have small crests and be unattractive to females, so you are
unlikely to get many grandchildren. Once it has become fashionable
to prefer big crests, genes that m a k e females defy the fashion are likely
to be eliminated.
Why should the fashion arise in the first place? I am going to tell
quite a complicated story involving several of a female's relatives, and
it will be easier to avoid confusion if she has a n a m e (Dinah). Dinah
has genes that m a k e her prefer big crests, and she chooses a big-crested
mate (Henry). Crest size is controlled by genes so it is likely that Hen-
ry's father also had a big crest. If he did, it is likely that Henry's m o t h e r
preferred big crests (otherwise she would probably not have chosen him).
This makes it likely that Henry has genes which, if he were female,
would make h i m prefer big-crested males. D i n a h ' s children are likely
to inherit genes for preferring big crests not only from Dinah, but also
from Henry. Futhermore, D i n a h ' s m o t h e r probably preferred big crests
so Dinah's father probably had one, and Henry's children will probably
inherit genes for big crests from Dinah as well as from h i m . These
effects are likely to m a k e crests bigger and preferences stronger in suc-
cessive generations. M a t h e m a t i c a l analysis shows that this kind of
evolution is likely to gather m o m e n t u m and go to extremes.
One of the things that m a k e s it seem likely that hadrosaur crests
evolved by sexual selection is that their shapes are so different in dif-
ferent species. If they had a function other than fashion they would
tend to have the same shape, the best shape for the job.
Figure 6.8 shows a Parasaurolophus crest sectioned to reveal the cav-
ities inside. You can see how the nasal passages start on the snout, run
all the way to the tip of the crest and double back to reach the m o u t h
cavity. This is a long route from nostrils to m o u t h even in this short-
crested (presumably female) skull. It is about t w o meters (6V2 ft) in the
long-crested (presumably male) Parasaurolophus shown in figure 6.6.
Tubes can be used to produce musical sounds, as in organs, flutes,
and trumpets. They can be made to resonate at particular frequencies,
emitting the regularly repeating waves of sound that we recognize as
86 FIGHTING A N D SINGING DINOSAURS
FIGURE 6 . 8 . T h e c r e s t of a Parasaurolophus s k u l l s e c t i o n e d t o s h o w t h e n a s a l
cavities. T h i s is probably a female skull, and it is not the s a m e species as the
Parasaurolophus i n f i g u r e 6 . 6 . F r o m H o p s o n 1 9 7 5 .
musical. We also use the resonant properties of our nasal passages in

voice production. It seems possible that Parasaurolophus used the long
tubes in its crest to produce sounds. T h e males may have done this to
attract females.
T h e resonant frequencies of air-filled tubes depend on their lengths.
Long organ pipes give lower notes than short ones, and you get the
lowest notes from a trombone by extending it to m a x i m u m length. I
will try to explain why, using a different resonant system to introduce
a basic principle.
Imagine a ball suspended on the end of a thin strand of rubber (figure
6.9). You can m a k e it vibrate up and down by moving your hand, but
there is one particular frequency of m o v e m e n t at which very small
hand m o v e m e n t s will sustain very large vibrations of the ball. That is
the resonant frequency. It exists because of interaction between the
mass of the ball and the elastic compliance of the rubber. "Compli-
a n c e " may be an unfamiliar word. It m e a n s stretchiness, in this sense:
if a force of one n e w t o n stretches the rubber by a centimeter, two new-
tons stretch it t w o centimeters, and so on, the compliance is one cen-
timeter per newton. You can reduce the resonant frequency either by
increasing the mass (using a heavier ball) or by increasing the compli-
ance (using a longer or thinner piece of rubber).
T h e air-filled cavity in figure 6.9b resonates in essentially the same
way. The vibrating mass is the air in the neck which vibrates in and
out, compressing the air inside, which has elastic compliance. T h e air
in the neck behaves like the ball and the air in the m a i n cavity behaves
like the strand of rubber but the distinction between the two lots of
air is of course blurred. The distinction is even more blurred in figure
6.9c, which shows air vibrating in a tube that is closed at one end. T h e
air near the m o u t h of the tube functions as a vibrating mass and the
air near the closed end as an elastic compliance, but there is a gradation
between the two. However, it is clear that a long tube contains a greater
mass of air than a shorter tube of the same diameter. It also has higher
compliance, because a long c o l u m n of air is squeezed up more, by a
given force on its end, than a short c o l u m n would be. T h e air in a
longer tube has more mass and compliance, so its resonant frequency
is lower.
Figure 6.9d shows what happens in a tube open at both ends. T h e
air at the ends vibrates in and out, compressing the air in the middle
and allowing it to expand again. T h e resonant frequency is the same
as for a tube with one closed end, of half the length. T h e air in a clar-
inet vibrates as in figure 6.9c but the air in a flute vibrates as in figure
6.9d because of the big hole at the blowing end.
The hollow crest of Parasaurolophus would probably have resonated
as a tube open at both ends. T h e resonant frequency of such a tube is
given approximately by a simple rule:
frequency in cycles per second = 170 + length in meters
The length of the nasal passages in the crest of the Parasaurolophus

in figure 6.6c is two meters, and the U-bend in t h e m m a k e s no differ-
F I G U R E 6.9. D i a g r a m s o f r e s o n a n t s y s t e m s : (a) a b a l l s u s p e n d e d b y a s t r a n d
o f r u b b e r ; |b) a n a i r - f i l l e d c a v i t y ; (c) a t u b e w i t h o n e e n d c l o s e d ; a n d (d) a t u b e
open at both ends.
FIGHTING AND SINGING DINOSAURS
ence to the calculations. Their resonant frequency must have been about
170/2 = 85 cycles per second, close to the note
Females with shorter crests (figure 6.8) would have had higher-pitched
voices. T h e equation m a k e s it possible to estimate the pitch of the
dinosaur's voice but tells us nothing about the tone, whether it was a
pure single-frequency sound like a note from a French horn or enriched
by m a n y harmonics like a note from a bassoon.
A big dinosaur should have a deafening roar, or so you may think.
However, there is a problem. You need a big loudspeaker to play a low
note loudly, and t r u m p e t s would not be so loud if they were not flared
out at the ends. T h e general rule is that to be most effective, loud-
speakers and the m o u t h s of t r u m p e t s should have diameters of at least
one-sixth of the wavelength of the sound. T h e wavelength is the speed
of sound (330 meters per second) divided by the frequency, so it is large
for low frequency sounds. For a frequency of 85 cycles per second it is
four meters, so an ideal loudspeaker for that pitch would have a di-
a m e t e r of almost 70 centimeters (28 in). Parasaurolophus should ide-
ally have had nostrils flaring out like the m o u t h s of t r u m p e t s 70 cen-
timeters in diameter. They should have been at least as big as the m o u t h s
of tubas designed to play similar low notes. There is no sign of their
being like that, so the dinosaur may not have been terribly noisy (if it
sang at all): it m a y have been more like a bassoon (which plays low
notes rather quietly) than a tuba.
Parasaurolophus had a long crest w i t h a tube inside but the other
hadrosaurs in figure 6.6 had shorter crests w i t h bulbous cavities that
would have resonated like the chamber in figure 6.9b. They too could
have been used to produce sound.
Whether the resonators were tubular or bulbous, bigger ones would
resonate at lower frequencies: larger crests would give deeper voices.
If females preferred large crests they probably also evolved a preference
for the deeper notes they produced. The hadrosaurs with the bass voices
got the females.
T h e message of this chapter is that in m a n y animal species, males
compete for females. It seems likely that male ceratopians fought for
females by wrestling with their horns like stags, and that pachyce-
phalosaurs fought by butting like rams. T h e crested hadrosaurs, how-
ever, probably completed more peaceably, letting the females choose
t h e m . T h e females had evolved a preference for large crests and deep
voices, and chose males partly for those qualities.
Principal Sources
C l u t t o n - B r o c k (1982) t e l l s a b o u t stags fighting a n d G e i s t (1971) t e l l s a b o u t r a m s .

T h e d i m e n s i o n s o f a n t e l o p e h o r n s i n figure 6.3 c o m e from P a c k e r (1983) a n d t h o s e
of Triceratops from H a t c h e r (1907). F a r l o w a n d D o d s o n (1975) d i s c u s s e d t h e h o r n s
and frills of c e r a t o p i a n s a n d S u e s (1978) d e s c r i b e d t h e s k u l l roof of p a c h y c e p h a l o -
s a u r s . A n d e r s s o n (1982) did t h e e x p e r i m e n t s o n w i d o w b i r d t a i l s a n d D a w k i n s (1986)
h a s w r i t t e n a good a c c o u n t of sexual selection. H o p s o n (1975) and W e i s h a m p e l (1981)
described a n d d i s c u s s e d t h e c r e s t s of h a d r o s a u r s .
A n d e r s s o n , M. 1982. F e m a l e c h o i c e s e l e c t s for e x t r e m e tail l e n g t h in a w i d o w b i r d .

Nature 2 9 9 : 8 1 8 - 2 0 .
Brown, B. a n d E. M. Schlaikjer, 1937. T h e s k e l e t o n of Styracosaurus w i t h t h e de-
s c r i p t i o n of a n e w s p e c i e s . American Museum Novitates 9 5 5 : 1 - 1 2 .
C l u t t o n - B r o c k , T. H. 1982. T h e f u n c t i o n s of a n t l e r s . Behaviour 7 9 : 1 0 8 - 1 2 5 .
D a w k i n s , R. 1986. The Blind Watchmaker. L o n d o n : L o n g m a n .
Farlow, J. O. a n d P. D o d s o n , 1975. T h e b e h a v i o u r a l s i g n i f i c a n c e of t h e frill a n d h o r n
m o r p h o l o g y in c e r a t o p s i a n d i n o s a u r s . Evolution 2 9 : 3 5 3 - 3 6 1 .
G e i s t , V. 1971. Mountain Sheep. C h i c a g o : U n i v e r s i t y of C h i c a g o Press.
Hatcher, J. B. 1907. T h e Ceratopsia. Monographs of the U.S. Geological Survey 4 9 : 1 -
300.
H o p s o n , J. A. 1975. T h e e v o l u t i o n of c r a n i a l d i s p l a y s t r u c t u r e s in h a d r o s a u r i a n di-
nosaurs. Paleobiology 1:21-43.
Packer, C. 1983. S e x u a l d i m o r p h i s m : T h e h o r n s of African a n t e l o p e s . Science
221:1191-1193.
Sues, H . - D . 1978. F u n c t i o n a l m o r p h o l o g y o f t h e d o m e i n p a c h y c e p h a l o s a u r i d di-
nosaurs. Neues fahrbuch fiir Geologie und Paldontologie, Monatshefte 1978(8):459-
472.
W e i s h a m p e l , D. B. 1 9 8 1 . T h e n a s a l c a v i t y of l a m b e o s a u r i n e h a d r o s a u r i d s . journal
of Palaeontology 55:1046-1057.
VII
Hot-Blooded Dinosaurs?
P EOPLE M A K E a big distinction between warm-blooded and cold-
blooded animals. T h e warm-blooded ones are the m a m m a l s and

birds, which feel w a r m and (if not too fierce) are nice to cuddle. Their
bodies are wrapped in heat-insulating fur or feathers. T h e cold-blooded
ones are snakes (and other reptiles), frogs (and other amphibians), fishes,
and all the invertebrates. They usually feel cold w h e n you touch them,
and m a n y of t h e m have scaly or slimy skin. Were the dinosaurs warm-
blooded or cold-blooded?
I w a n t to rephrase the question. Scientists do not like talking about
"warm-blooded" and "cold-blooded" animals because the "cold-blooded"
ones are not necessarily colder than the "warm-blooded." Birds keep
their bodies at 4 0 - 4 3 ° C and m a m m a l s at 36-40°C, in all climates. Rep-
tiles, amphibians, and fishes have more varied temperatures that de-
pend on their surroundings. Most fishes have temperatures almost ex-
actly the same as the water they are living in, w h e t h e r it be an arctic
lake or a tropical swamp. Many reptiles, however, adjust their tem-
peratures on sunny days by moving back and forth between sun and
shade. If they stayed in the shade all day they would be rather cool
and if they stayed in tropical sun all day they would get dangerously
hot, but by moving in and out of the sun some manage to keep their
temperatures in the m a m m a l range for most of the day. Should we call
t h e m cold-blooded?
We scientists prefer to talk about e c t o t h e r m s and endotherms. T h e
ectotherms are the "cold-blooded" animals. If they control their body
temperatures they do so mainly by using heat from the sun. T h e en-
dotherms however depend mainly on heat produced within their bod-
ies, by the chemical processes of metabolism. The best-known en-
dotherms are birds and m a m m a l s but I will m e n t i o n some mildly
endothermic fish in chapter 9.
HOT-BLOODED DINOSAURS? 9 1
Metabolism supplies the energy needed to keep the body working.

It happens in all living things, but particularly fast in endotherms. T h e
most important processes of m e t a b o l i s m combine food w i t h oxygen to
give (mainly) carbon dioxide and water, and release energy. T h e nec-
essary oxygen is obtained by breathing and the carbon dioxide is got
rid of in the breath. The basic process is the same as burning, and the
quantity of energy released is the same as if the food had been burnt
to give the same end products. Some of the energy does useful jobs,
powering muscles and driving the m a n y energy-absorbing processes of
life, but these processes are rather inefficient, and most of the energy
is released as heat. The heat that w a r m s our bodies is a waste product
of metabolism.
The bodies of animals work by chemical processes, and chemical
processes generally go faster at higher temperatures. High body tem-
peratures enable animals to run faster than they otherwise could, to
digest food faster, and (if food is abundant), to grow faster. For example,
the lizard Iguana can run nearly three times as fast w h e n its body tem-
perature is 35°C, as when it is 20°C. Too high a temperature would be
lethal but within limits it is good to be warm.
Metabolism goes fastest w h e n animals are active but does not stop
when they sit still. Even w h e n resting they need energy to keep the
heart beating and for m a n y less obvious processes that are essential for
life. When an animal is inactive its metabolism falls to a low, resting
rate, which depends on the temperature. Ectotherms have higher rest-
ing metabolic rates at higher temperatures. Endotherms, however, in-
crease their metabolic rates w h e n they are put in cold places, to pro-
duce the heat needed to m a i n t a i n their body temperatures.
The metabolic rates of m a m m a l s can generally be measured, by mea-
suring how fast they use oxygen. This is done by putting the animal
in a sealed container and analyzing samples of the air from t i m e to
time to find out how m u c h of its oxygen has been used. (Obviously,
the oxygen concentration m u s t not be allowed to fall too m u c h . Oth-
erwise the animal's metabolism would be affected and it would even-
tually suffocate.) Whatever food is being metabolized, about 20 joules
of energy are released for every cubic centimetre of oxygen used.
Figure 7.1 shows resting metabolic rates plotted against body mass,
for lizards, birds, and m a m m a l s . It includes data for animals of a wide
range of sizes, from sparrows to elephants. To show all these data clearly
I have distorted the scales of the graph in the same way as I distorted
the graph of bone circumference against body m a s s (figure 2.5): I have
made the scales proportional to the logarithms of the quantities they
represent so that the distance from 1 to 10 units is the same as from
10 to 100 units, or from 100 to 1,000 units.
9 2 HOT-BLOODED DINOSAURS?
T h e metabolic rates shown for lizards in figure 7.1 are resting rates
at 37°C. T h e lizards could not have stood m u c h higher temperatures,
so these are n e a r - m a x i m u m resting metabolic rates. T h e rates shown
for m a m m a l s and birds, however, are m i n i m u m rates measured at
comfortable environmental temperatures, at which the metabolism
needed for other essential processes is enough to maintain the tem-
perature of the body. T h e rates for m a m m a l s are nevertheless five to
eight times as high as for lizards of the same mass, and some birds
have even higher rates. M a m m a l s and birds are enabled to be endo-
thermic by their high metabolic rates. Even w h e n they are resting at
comfortable temperatures their metabolism produces heat fast enough
to w a r m t h e m quite a lot.
Dinosaurs are usually classed as reptiles. They are closely related to
the crocodiles which, like other modern reptiles, are ectotherms. How-
ever, they are also closely related to the endothermic birds. Dr. Robert
HOT-BLOODED DINOSAURS? 93
Bakker argues that dinosaurs were e n d o t h e r m s . Some of his arguments

are not (to me) very convincing. He points out that dinosaurs moved
like m a m m a l s and birds, w i t h their feet under the body (figure 3.5) and
that some of the smaller ones were lightly built like ostriches. This
suggests to h i m that dinosaurs, like ostriches, were fast-moving, fast-
metabolizing endotherms. He also points out that the microscopic
structure of dinosaur bone is like that of birds and m a m m a l s , which
suggests to h i m that dinosaurs, like birds and m a m m a l s , were endo-
therms. (The microscopic structure can be seen in polished slices of
fossil bone.)
These kinds of evidence are indirect. There is no clear reason why
ostrich-like proportions or mammal-like bone structure should be found
only in endotherms. However, Bakker presented a third argument w i t h
a m u c h clearer logical basis. Endotherms metabolize m u c h faster than
ectotherms, so they have to eat a great deal more. An e n d o t h e r m i c
predator needs a bigger herd of prey to keep it supplied with food, than
an ectotherm would do. If we can estimate the relative population sizes
of carnivorous dinosaurs and their prey, we may discover w h e t h e r the
dinosaurs could have been endotherms. A population of endothermic
carnivores would need a m u c h bigger population of prey to support
them, than would a similar-sized ectothermic population.
There are two difficulties about this line of argument. T h e first is
that it is difficult to be sure just how big a population of prey is needed
to feed a population of endothermic predators. Obviously reproduction
and growth of the prey population m u s t be enough to compensate for
what the predators eat (otherwise the prey population will dwindle and
the predators will starve) but it is hard to calculate how big a prey
population that needs.
Ecologists have counted the large m a m m a l s in m a n y nature reserves
in Africa, India and America. They have estimated, for example, that
each square kilometer of the Ngorongoro crater in Tanzania supports
10.4 tonnes of herbivores (antelopes and zebra) but only 96 kilograms
of carnivores (lions and hyaenas): the mass of carnivores is only 1 per-
cent of the mass of herbivores. In 29 other reserves the carnivore m a s s
was always less than 2 percent of the herbivore mass. Bigger carnivore
populations probably could not find enough food.
In contrast, in populations of ectothermic predators and prey, the
predator mass may total 10 to 40 percent of the prey mass. This has
been found, for example, for populations of spiders and ants feeding on
insects. We would like to have data for c o m m u n i t i e s in which both
the principal predators and the principal prey were reptiles, but, un-
fortunately, there do not seem to be any modern communities like that.
However, it does seem clear that, in a c o m m u n i t y of ecothermic di-
9 4 HOT-BLOODED DINOSAURS'
nosaurs, the predator mass could be far more than 2 percent of the prey
mass.
T h e second difficulty about deciding whether dinosaurs could have
been endotherms, by measuring the ratio of predators to prey, is that
the ratio is very difficult to estimate for extinct animals. The best di-
nosaurs to try w i t h (because there are lots of them) seem to be the ones
that have been found in Canada in some late Cretaceous rocks, the
O l d m a n Formation. A count of these dinosaurs gave 246 individual
herbivores (mainly ornithopods) estimated to have an average mass of
5 tonnes, and 22 carnivores (tyrannosaurs) averaging 2 tonnes. This
m a k e s 1,230 tonnes of herbivores to 44 tonnes of carnivores. However,
the c o m m o n e r herbivores in the O l d m a n Formation are so very com-
mon that the paleontologists who collected there probably did not bother
to collect the less good specimens. An informed guess about how m a n y
they left increases the estimated mass of herbivores to 2,110 tonnes.
This m a k e s the carnivore mass 2 percent of the herbivore mass.
If the dinosaurs had all been killed at once by a volcanic eruption or
some other calamity, we could conclude that the carnivore population
had had 2 percent of the m a s s of the herbivore population. There is no
sign of such a calamity and we m u s t suppose that the dinosaurs died
naturally of various causes. In that case the n u m b e r s of fossils probably
do not reflect the sizes of the living populations, but the rates at which
they died. T h e herbivores probably had shorter lives than the carni-
vores, because they were liable to be attacked and eaten, so there is
probably a bigger proportion of herbivores among the fossils than in
the living population. The carnivore population probably had more than
2 percent of the mass of the herbivore population, which makes it
doubtful w h e t h e r they could have had m a m m a l - l i k e metabolism.
We might conclude that the dinosaurs had reptile-like metabolism,
or at least that their m e t a b o l i s m was slower than would be expected
for m a m m a l s of equal size. However, I would be wary of reaching any
firm conclusion on this evidence. The conclusion could easily be changed
if it turned out that paleontologists had discarded even more of the
less-good herbivore fossils than has been supposed.
O n e argument that m i g h t be used against the suggestion that di-
nosaurs were endotherms is that they had neither fur nor feathers. Most
m a m m a l s have fur and birds have feathers to retain the heat produced
by metabolism, but the few fragments of dinosaur skin that have been
preserved as impressions in rocks are bare and scaly like the skin of
modern reptiles. However, these fragments are from the skin of large
dinosaurs, and very large modern m a m m a l s (elephants, rhinoceros, hip-
popotamus) also have no fur.
Endothermy is a m a t t e r of degree. A lizard might be able to metab-
olize fast enough to keep it a degree or two warmer than its environ-
ment, but that would not m a k e you want to call it an endotherm: it
could warm itself far more by the e c t o t h e r m i c m e t h o d of basking in
the sun. We would not call dinosaurs e n d o t h e r m s unless their metab-
olism could keep t h e m a good deal w a r m e r than their environments.
Could they have done this, in spite of their having no fur? If they
were warmer than their surroundings they would lose heat, and to keep
a constant temperature they would have to produce heat fast enough
by metabolism to replace the losses. I will try to calculate how w a r m
they could have kept themselves both if they had reptile-like metab-
olism and if they had faster, m a m m a l - l i k e metabolism. I will base the
calculations on the rates at which modern reptiles cool, w h e n moved
from a warm place to a cooler one, so we will need to k n o w s o m e t h i n g
about the physics of cooling.
To show how things cool, I baked a potato in its jacket in the oven.
When it had cooked I pushed a t h e r m o m e t e r into it and left it to cool
on the kitchen table. Figure 7.2 shows h o w its temperature fell. Ini-
tially it was 93°C, which was 72° above room temperature. After 40
m i n u t e s it was about 36° above room temperature, after 80 m i n u t e s it
was about 18° above room temperature and after 120 m i n u t e s it was
only about 9° above room temperature: the temperature difference halved
every 40 m i n u t e s . Hot objects in constant e n v i r o n m e n t s generally cool
F I G U R E 7.2. A g r a p h o f t e m p e r a t u r e a g a i n s t t i m e for a c o o l i n g p o t a t o .
96 HOT-BLOODED DINOSAURS?
like this, taking equal times for each successive halving of the tem-
perature difference (figure 7.3a). This is called exponential cooling.
Obviously, some things cool faster than others. We could use the
half t i m e (the t i m e the temperature difference takes to halve) as a mea-
sure of the rate of cooling. Instead, I am going to do something that
may seem capricious. I am going to use the t i m e taken for the differ-
ence to fall to 0.37 of its initial value. This t i m e is called the cooling
time constant.
There is something very special about that number, 0.37. It is the
reciprocal of the n u m b e r called e, the base of natural logarithms, which
is the second most useful n u m b e r in the whole of m a t h e m a t i c s . (The
most useful is the one called pi). Figure 7.3b shows w h y it is useful to
us here. T h e curve is exactly the same as in figure 7.3a. T h e temper-
ature difference starts at D and reaches 0.37D after t i m e T, so T is the
t i m e constant. Cooling starts fast and gradually gets slower, but the
sloping broken line shows that if it had continued at its initial rate the
temperature difference would have reached zero at time T. In other
words the cooling rate was D/T when the temperature difference
was D.
D/T is the rate of fall of temperature, but we want to calculate the
rate of loss of heat. We do this by multiplying by the heat capacity C,
the a m o u n t of heat energy that m u s t be gained or lost to change the
temperature of the object by one degree. T h u s the rate of heat loss from
an object, when its temperature is D above its surroundings, is CD/T.
We want to k n o w how m u c h dinosaurs would have been heated by
their own metabolism. To keep body temperature constant, their rates
of heat loss (CD/T) would have to be matched by their metabolic rates
[R): R = CD/T. D is the temperature difference between the dinosaur
and its environment, the quantity that we want to know. We get it by
rearranging the equation D = RT/C.
We will use figure 7.1 to estimate R for different-sized dinosaurs with
reptile-like or m a m m a l - l i k e metabolism. C is easy to estimate for a
dinosaur of any particular mass because the heat capacity of animal
tissue is about 3,500 joules per kilogram, a little less than for pure
water. Finally, we will get values for T from the results of experiments
on modern reptiles.
In these experiments, living reptiles were put somewhere warm un-
til their body temperatures (measured by a tiny electrical t h e r m o m e t e r
in the gut) had risen to 35 or 40°C. They were then moved to a cooler
place and their temperatures were recorded as they cooled. Their met-
abolic rates were too low to have a noticeable effect on their temper-
atures so they cooled exponentially, like the hot potato. Big ones cooled
HOT-BLOODED DINOSAURS? 9 7
F I G U R E 7 . 3 . (a) A g r a p h o f t e m p e r a t u r e d i f f e r e n c e a g a i n s t t i m e , s h o w i n g t h e
difference h a l v i n g in s u c c e s s i v e e q u a l i n t e r v a l s of t i m e . D is t h e initial t e m -
p e r a t u r e d i f f e r e n c e a n d t i s t h e t i m e i n w h i c h i t h a l v e s , (b) T h e s a m e g r a p h ,
s h o w i n g h o w the t i m e constant T is defined.
more slowly than small ones just as big potatoes cool more slowly than
small potatoes.
In some of the experiments, alligators and other semi-aquatic rep-
tiles were cooled in water. Each reptile cooled faster in water than in
air at the same temperature. This is what you might expect, but we
need to understand why. T h e reptile's body can be thought of as a cen-
tral core, kept at a uniform temperature by the circulating blood, and
an outer heat-insulating layer of skin (figure 7.4a). Heat loss is a two-
stage process. First the heat is conducted through the skin and then it
leaves the skin surface by convection and radiation. Convection means
the process of heat being carried away from the surface by moving air
or water. These m o v e m e n t s may be convection currents due to hot
fluid rising or winds or water currents due to other causes. Convection
works m u c h more effectively in water than in air because water has a
m u c h higher heat capacity than an equal volume of air. T h i s is why
reptiles cool faster in water.
Figure 7.4b shows cooling t i m e constants for a range of sizes of rep-
tiles. T h e lines for cooling in air and in water converge, as body mass
increases, suggesting that reptiles of 100 kilograms or more would cool

almost as fast in air as in water. For them, heat loss would be limited
almost entirely by the insulating effect of the skin. Theory predicts
that the graphs should converge, when plotted on logarithmic scales
as in figure 7.4b. T h e graph for cooling in air should be curved and the
graph for cooling in water straight, which is h o w the graphs seem
to be.
In other experiments reptiles were heated instead of being cooled:
they were put in a cool place until their body temperatures had dropped
to 15° to 20°C, and then moved to a w a r m one. Their temperatures
were recorded as they warmed up and t i m e constants were calculated.
These time constants were shorter than the t i m e constants for cooling,
usually about half as long. I have included the t i m e constants for
warming in water in figure 7.4b but not those for warming in air, to
avoid confusing the graph. If I had included the line for warming in air
it would have run below and roughly parallel to the line for cooling
in air.
Here is a likely explanation for the difference between cooling and
warming. The reptiles probably let more blood flow to their skins when
they were warming, reducing the effective thickness of the insulating
layer of skin. They were being warmed from low temperatures to their
preferred temperatures, so would w a n t to w a r m as quickly as possible.
Conversely, in the cooling experiments they were probably delaying
cooling by restricting blood flow to the skin, m a k i n g the insulating
layer as thick as possible. People similarly adjust blood flow to their
skin to help control body temperature: that is why we get flushed when
we are overheated.
Dinosaur skin seems to have been like the skin of modern reptiles,
so it seems reasonable to estimate t i m e constants for dinosaurs by ex-
tending the graphs to higher body masses (figure 7.4b, broken lines).
The difference between the warming and cooling graphs shows the likely
scope for adjustment by controlling blood flow to the skin. T h e
straightness of the lines for warming and cooling in water m a k e s it
seem fairly obvious how the extensions should go, but there is a danger
of serious error because we have to extend the lines so far. Some di-
nosaurs were over 1000 times heavier than the biggest reptiles used in
the experiments. It seems better to risk error than to give up in despair.
T h e graph suggests that a 5-tonne Iguanodon would have had t i m e
constants of 2 - 4 days and a 50-tonne Biachiosauius 1-3 weeks, de-
pending on how m u c h blood they let flow to their skins.
This graph gives us the t i m e constants we need to estimate tem-
perature differences between dinosaurs and their e n v i r o n m e n t s . In ta-
ble 7.1 the "reptile-like" values a s s u m e that the dinosaur metabolizes
at the rate indicated by figure 7.1 for a reptile of its mass, resting with
its body at a temperature of 37°C. T h e " m a m m a l - l i k e " values assume
m e t a b o l i s m at the rate indicated for a m a m m a l resting in a comfort-
able w a r m e n v i r o n m e n t in w h i c h the metabolism necessary for other
purposes is enough to m a i n t a i n body temperature. The range of tem-
perature difference given in each case is from the value calculated from
the warming t i m e constant to the one calculated from the cooling t i m e
constant. It is the range that the animal could probably achieve by ad-
justing blood flow to the skin. Higher temperature differences would
be possible if the animals increased their metabolic rates, as modern
birds and m a m m a l s do in cold conditions. In experiments with Car-
dinal finches the birds doubled their resting metabolic rates when moved
from an e n v i r o n m e n t at 15°C to one at - 1 5 ° C . Lower temperature dif-
ferences than those s h o w n in the table would also be possible if the
animals panted to increase the a m o u n t of water lost by evaporation
into their breath. T h e heat needed to evaporate the water would be
lost from the body.
The table shows larger temperature differences for big dinosaurs than
for small ones w i t h the same type of metabolism, because bigger di-
nosaurs have longer t i m e constants. It also (obviously) shows larger
temperature differences for dinosaurs w i t h fast, m a m m a l - l i k e metab-
olism than for ones with slow, reptile-like metabolism. It indicates that
50- and even 500-kilogram dinosaurs with reptile-like metabolism would
be very little w a r m e r than their surroundings: they would not be ef-
fective endotherms. Fifty-tonne dinosaurs with reptile-like metabolism
would be quite good endotherms, up to 13° warmer than their sur-
roundings. Moderate-sized dinosaurs with m a m m a l - l i k e metabolism
would be endotherms, despite their lack of fur or feathers, and 50-tonne
dinosaurs w i t h m a m m a l - l i k e m e t a b o l i s m would be in danger of getting
so hot that they cooked themselves, in most climates.
Small dinosaurs could not have been endotherms, even with m a m -
mal-like metabolism. R e m e m b e r the baby Psittacosaurus in figure 1.2,
smaller t h a n a pigeon. It is m u c h too small to have been an effective
endotherm unless (which seems unlikely) it had fur or feathers. Some
other baby dinosaurs were bigger, even w h e n newly hatched, but it
seems doubtful w h e t h e r any were big enough to be endotherms from
the start. Fossil dinosaur eggs are big, but they are not enormous. Even
some thought to have been laid by a sauropod are only 25 centimetres
long.
You would be wrong to assume that, because I have used scientific
arguments, I have got everything right. I have made rough calculations
involving extrapolation from the small reptiles used in experiments to
the big dinosaurs. I will try to check whether the results are plausible
by making some comparisons w i t h modern animals.
Table 7.1 indicates that 50- and even 500-kilogram dinosaurs w i t h
reptile-like metabolism would be effectively ectothermic. This fits what
we know of modern reptiles, all of w h i c h seem to be ectotherms. Even
large crocodiles are ectotherms.
No modern reptiles have m a m m a l - l i k e metabolism but it seems rea-
sonable to guess that m a m m a l s that have no fur have time constants
about equal to those of similar-sized reptiles. Most of these m a m m a l s
are large but the Naked mole rat [Heterocephalus) is small. It lives in
tropical Africa in underground burrows in w h i c h the temperature re-
mains very constant at about 30°C. Its body temperature is always close
to burrow temperature. It is in effect an ectotherm, as table 7.1 sug-
gests it would have to be. We ourselves are naked m a m m a l s with masses
of about 50 (women) or 70 kilograms (men). W i t h o u t clothes we are
comfortable only in w a r m climates, which seems consistent w i t h the
calculation in the table that 50-kilogram dinosaurs w i t h m a m m a l - l i k e
TABLE 7.1 Calculated temperature differences between dinosaurs

and their environments. These are equilibrium values for dinosaurs
resting in constant environments, taking no special steps to control
body temperature. Cooling by evaporation of water is ignored.
Body mass 5 0 kg 5 0 0 kg 5 tonnes 50 tonnes
M e t a b o l i c r a t e ( w a t t s ) for
reptile-like m e t a b o l i s m 17 110 730 4,900
mammal-like metabolism 75 430 2500 14,000
T i m e c o n s t a n t (days) f r o m
cooling experiments 0.13 0.69 3.7 20
warming experiments 0.07 0.34 1.6 8
H e a t c a p a c i t y ( m e g a j o u l e s p e r °C) 0.18 1.8 18 180
T e m p e r a t u r e d i f f e r e n c e (°C) for
reptile-like m e t a b o l i s m 0.2-0.4 0.7-1.4 2-4 6-13
mammal-like metabolism 3-5 7-15 20-50 60-140
T h e calculated temperature difference |°C| is:

m e t a b o l i c rate (watts) x t i m e c o n s t a n t (seconds)
heat capacity (joules per °C)

metabolism could keep themselves only a few degrees warmer than

their surroundings. Elephants seem to have trouble keeping cool in Af-
rica, and seem comfortable enough even in winter in zoos in temper-
ature countries, which seem consistent w i t h the entry in the table for
5-tonne dinosaurs.
Those comparisons encourage me to think that the calculations may
have been reasonably realistic, but another comparison worried me at
first. The table says that a 50-tonne dinosaur w i t h mammal-like me-
tabolism would be at least 60° w a r m e r than its surroundings unless it
did something to get rid of excess heat. However, 50-tonne whales must
always be less than 40° warmer than the water, even in polar seas.
Remember that such large animals lose heat little faster in water than
they would in air at the same temperature. If large whales are possible,
should not equally large dinosaurs w i t h m a m m a l - l i k e metabolism also
be possible?
One fault of my argument so far is that I have almost ignored loss
of heat by evaporation of water. I m e n t i o n e d panting but have not con-
sidered the water loss that must occur all the time. The insides of lungs
are damp, so water evaporates from t h e m and gets lost in the breath.
Skin is slightly permeable to water, so some water diffuses out through
it and evaporates: crocodiles lose a lot of water this way, but desert
lizards, w i t h less permeable skins, lose only a little. I have written as
if all heat loss from the bodies of reptiles happened by conduction
through the skin followed by convection and radiation from the skin
surface. We m u s t r e m e m b e r that heat is also removed by evaporating
water.
Suppose that a 50-tonne dinosaur were living in a warm climate,
where the temperature of its surroundings was the same as its preferred
body temperature. To keep its body at that temperature it would have
to lose water fast enough to remove all the heat produced by its me-
tabolism. The heat needed to evaporate one gram of water is 2,500 joules
so if it had m a m m a l - l i k e metabolism, producing heat at the rate of
14,000 w a t t s (joules per second; see table 7.1) it would have to evap-
orate about six grams of water per second, or about half a tonne (1
percent of its body mass) per day. A 5-tonne dinosaur with m a m m a l -
like metabolism, in similar circumstances, would have to lose about
90 kilograms of water (1.8 percent of body mass) per day. These rates
do not seem impossible. A large (3.7 tonne) Indian elephant drank 140
kilograms of water per day and lost about 20 kilograms of it (0.5 per-
cent of body mass) by evaporation. (The rest was lost in urine and feces.)
This was w h e n it was kept at a comfortable temperature, 20°C: in hot
conditions it could presumably have allowed more to evaporate, and if
necessary drunk more. Large dinosaurs w i t h m a m m a l - l i k e metabolism
need not have overheated, even in quite w a r m climates, if they allowed

enough water to evaporate from their skin and into their breath.
I hope that argument has convinced you that you should not take
the temperature differences in table 6.1 too literally. They could prob-
ably all be reduced to zero if the a n i m a l s allowed enough water to evap-
orate from their bodies. However, they need not be far wrong for an-
imals trying to conserve water: animals can, if necessary, lose a lot of
water by evaporation and they m u s t inevitably lose some (unless the
atmosphere is very humid), but they need not lose m u c h . T h e evapo-
ration from the elephant at 20°C dissipated only 20 percent of the heat
produced by its metabolism.
T h e ignoring of evaporation is not the only unrealistic thing about
table 7.1. The table assumes resting metabolic rates, but animals do
not rest all the time. Their metabolic rates are higher when they are
active, and even at rest e n d o t h e r m s can increase their metabolic rates
to maintain their body temperatures in cold conditions.
Despite these points the table has (I think) some value. Make any
reasonable allowances for activity and water loss, and it will still show
that only very big dinosaurs could have been effective endotherms, if
they had reptile-like metabolism.
I have argued that even with m a m m a l - l i k e metabolism, big dino-
saurs need not have overheated, if they allowed enough water to evap-
orate from their bodies. However, m a n y hot e n v i r o n m e n t s are also dry,
with water in short supply. It may be best to lose as m u c h excess heat
as possible by convection and radiation, keeping evaporation to a min-
i m u m . Elephants use their large ears as radiators and convectors. In
hot conditions they dilate the blood vessels of their ears, and flap the
ears to lose as m u c h heat as possible from the blood passing through.
It has been suggested that the plates on the backs of stegosaurs (figure
1.13) may also have served as cooling devices. Experiments with models
in a wind tunnel seemed to confirm that the idea is feasible.
I have written so far as if an animal's surroundings had a clearly
defined temperature. There is no problem for aquatic animals: for t h e m
the temperature of the water is the only external temperature that mat-
ters. A terrestrial animal, however, m a y be surrounded by air at one
temperature, ground at another and foliage at a third. T h e atmosphere
above it has different temperatures at different levels. T h e animal is
affected by all these temperatures because it exchanges heat w i t h the
ground, foliage and sky by radiation, as well as exchanging heat w i t h
the immediately surrounding air by convection. Also, if the sun is
shining its rays may have a profound effect on the animal's heat bal-
ance. There is, however, a temperature that we can think of as the
effective temperature of the animal's surroundings. It is the tempera-
104 HOT-BLOODED DINOSAURS'
ture at which the animal's body would eventually settle if it did not
metabolize or lose water by evaporation, and if conditions remained
constant.
In practice, conditions do not remain constant. It is warmer during
the day and colder at night. T h e body temperatures of modern ecto-
t h e r m s fluctuate accordingly. For example, an investigation of the liz-
ard Amphibolurus in Australia showed that its body cooled to 25°C
during s u m m e r nights but w a r m e d to almost 40°C during the day. T h e
temperatures of large dinosaurs m u s t have fluctuated far less, because
of their long t i m e constants. They would hardly have started heating
up during a hot day, before the cool night came. A 50-tonne Brachio-
saums, w i t h estimated t i m e constants of 8 to 20 days, m u s t have had
almost constant body temperature day and night. It may have got hot-
ter in s u m m e r and cooler in winter, but its daily temperature fluctua-
tions m u s t have been slight. Even if it had reptile-like metabolism it
could have maintained a high body temperature day and night, because
of its huge size.
We started with the question, were dinosaurs ectotherms or endo-
therms? I explained the t e r m s and showed that modern reptiles (which
are ectotherms) have m u c h slower metabolism than similar-sized birds
and m a m m a l s (which are endotherms). I explained why dinosaurs have
been thought to be e n d o t h e r m s . T h e most direct argument depended
on the relative c o m m o n n e s s of predatory dinosaurs and their prey.
I explained some of the basic physics of cooling and showed how
rates of heating and cooling of modern reptiles could be used to esti-
m a t e rates of heat loss from dinosaurs. The calculations seemed to show
that small dinosaurs w i t h reptile-like metabolic rates would be ec-
t o t h e r m s (like modern reptiles) but that very large ones would be quite
good e n d o t h e r m s . If dinosaurs had fast, m a m m a l - l i k e metabolism, very
small ones (including hatchlings) would nevertheless be ectotherms
unless they had fur or feathers. Moderate-sized ones would be endo-
t h e r m s and the largest would have been liable to overheat unless they
lost a lot of water by evaporation from their skin and in their breath.
Whether they had reptile-like or m a m m a l - l i k e metabolism, the body
temperatures of large dinosaurs would have remained almost constant,
day and night.
I have to admit after all this discussion that I do not know whether
dinosaurs were ectotherms or endotherms, and whether they had
m a m m a l - l i k e or reptile-like metabolic rates. Both questions remain
to puzzle us. I would however like to present one thought that I find
startling.
Suppose that the dinosaurs had reptile-like metabolism. Suppose also
that plants grew as lushly as they do now, so that there was as m u c h
for herbivorous dinosaurs to eat as there is now for herbivorous m a m -

mals. If both those things were true, large dinosaurs may have been
remarkably n u m e r o u s . Figure 7.1 indicates that large dinosaur-sized
reptiles would use energy at about the same rate as m a m m a l s of about
one-fifth their mass. Therefore they need only as m u c h food as m a m -
mals of one-fifth their mass. T h i n k of the parts of East Africa where
there are still large herds of m a m m a l s , including gazelles, wildebeest,
zebra, buffalo, and elephant. T h i n k of the population of dinosaurs that
such a place could support if its vegetation was as plentiful in Meso-
zoic times. (The vegetation would have been different from modern
vegetation but I see no reason why it should not have been as plentiful
and have grown as fast.) For every 500-kilogram buffalo that lives now
the vegetation could have supported a 2.5-tonne stegosaur, and for every
3-tonne elephant a 15-tonne Diplodocus. If dinosaurs had reptile-like
metabolism they may have been as n u m e r o u s as m a m m a l s of one fifth
their mass in modern populations. T h e world may have seemed very
full of dinosaurs.
Principal Sources
B a k k e r (1972) s u g g e s t e d t h a t t h e d i n o s a u r s w e r e e n d o t h e r m s a n d s t i m u l a t e d a great
deal of d i s c u s s i o n w h i c h he h a s r e v i e w e d in a r e c e n t b o o k (1986). F a r l o w (1976)
m a d e a m o r e t h o r o u g h i n v e s t i g a t i o n of t h e r a t i o of p r e d a t o r s to prey in a c o m m u n i t y
of large d i n o s a u r s . S o u r c e s for my d a t a on m e t a b o l i c r a t e s a n d on h e a t i n g a n d cool-
ing t i m e c o n s t a n t s c a n be f o u n d in C a l d e r (1984) a n d Bell (1980) r e s p e c t i v e l y . Spotila
et al. (1973) t o o k a different a p p r o a c h to d i n o s a u r h e a t b a l a n c e , m a k i n g a s s u m p t i o n s
a b o u t t h e t h i c k n e s s of t h e s k i n i n s t e a d of e x t r a p o l a t i n g from t i m e c o n s t a n t s of m o d -
e r n r e p t i l e s . Farlow et al. (1976) i n v e s t i g a t e d t h e p o s s i b i l i t y t h a t t h e p l a t e s of Ste-
gosaurus m a y h a v e b e e n c o o l i n g fins. 1 h a v e b e n e f i t e d g r e a t l y , in w r i t i n g t h i s c h a p -
ter, from h a v i n g s e e n s o m e m u c h m o r e e l a b o r a t e c a l c u l a t i o n s a b o u t d i n o s a u r h e a t
b a l a n c e , m a d e b y m y s t u d e n t (yrki H o k k a n e n .
Bakker, R. T. 1972. A n a t o m i c a l a n d e c o l o g i c a l e v i d e n c e of e n d o t h e r m y in d i n o s a u r s .
Nature 2 3 8 : 8 1 - 8 5 .
Bakker, R. T. 1986. The Dinosaur Heresies N e w Y o r k : M o r r o w .
Bell, C. J. 1980. T h e s c a l i n g of t h e t h e r m a l i n e r t i a of l i z a r d s , fournal of Experimental
Biology 8 6 : 7 9 - 8 5 .
Benedict, F. G. 1936. The Physiology of the Elephant. W a s h i n g t o n : C a r n e g i e Insti-
tution.
Calder, W. A. 1984. Size, Function, and Life History C a m b r i d g e M a s s . : H a r v a r d U n i -
v e r s i t y Press.
Farlow, ]. O. 1976. A c o n s i d e r a t i o n of t h e t r o p h i c d y n a m i c s of a late C r e t a c e o u s
l a r g e - d i n o s a u r c o m m u n i t y ( O l d h a m F o r m a t i o n ) . Ecology 5 7 : 8 4 1 - 8 5 7 .
Farlow, ]. O., C. V. T h o m p s o n , a n d D. E. R o s n e r . 1976. P l a t e s of t h e d i n o s a u r Ste-
gosaurus: Forced c o n v e c t i o n h e a t loss fins? Science 1 9 2 : 1 1 2 3 - 1 1 2 5 .
Spotila, J. R., P. W. L o m m e n , G. S. B a k k e n , a n d D. M. G a t e s . 1973. A m a t h e m a t i c a l
m o d e l for body t e m p e r a t u r e s of large r e p t i l e s : I m p l i c a t i o n s for d i n o s a u r e c o l o g y .
American Naturalist 107:391-404.
VIII
Flying Reptiles
T HE PTEROSAURS lived at the same time as the dinosaurs. Both groups

appeared late in the Triassic period and became extinct at the end
of the Cretaceous. Pterosaurs were winged reptiles, w i t h astonishingly
long fingers that helped to support their wings (figure 8.1). Most fossils
show only the skeleton, but a few show impressions of the wings and
one has marks that seem to be impressions of fur. If pterosaurs were
furry they were probably endotherms, w i t h the fur serving as heat in-
sulation. Flapping flight, as practised by birds and bats, is extremely
energetic. Being e n d o t h e r m s may have enabled pterosaurs to get the
necessary power o u t p u t from their muscles.
Describing pterosaurs as furry animals with wings makes them sound
pretty m u c h like bats, but they looked quite different. Bat wings con-
sist of a thin m e m b r a n e stretched between the body, the arm and four
extremely long fingers. Pterosaurs had only one finger, the huge "lit-
tle" one, involved in supporting the wing, which was m u c h more pointed
than bat wings. Some of the wing impressions that have been found
have striations on them, running in the same directions as the quills
of the m a i n wing feathers of birds (figure 8.1). If these are the remains
of stiffening rods, as some palaeontologists think, the wings may have
been relatively stiff structures like bird wings, not billowing m e m -
branes like the wings of bats and hang gliders.
Palaeontologists argue about the width of the wings. Figure 8.2a shows
on the left the traditional reconstruction, a wide wing attached, as in
bats, to the hind limbs as well as the fore. On the right it shows an
alternative reconstruction that has been supported very persuasively
by Dr. Kevin Padian of the University of California, Berkeley. This has
narrow wings attached to the t r u n k but not to the hind legs. T h e fos-
sils w i t h wing impressions seem to show that the wings were indeed
narrow.
FLYING REPTILES 107
FIGURE 8.1. Rhamphorhynchus, a p t e r o s a u r . T h e w i n g s p a n i s o n e m e t e r . F r o m

Wellnhofer 1975.
Figure 8.2b shows how supporters of the wide-wing hypothesis think

pterosaurs would have looked on the ground. T h e hind legs stick out
sideways like the legs of bats. T h e animal is imagined crawling awk-
wardly on all fours, hampered by the wing m e m b r a n e attached to all
four limbs. There is a famous set of fossil footprints that looks as if it
was made by a pterosaur crawling like this but Padian has shown (sur-
prizingly) that big lizards m a k e very similar footprints. T h e m a r k s that
had been interpreted as impressions of the base of the long finger are
imitated, in the lizard prints, by scratch made by a toe as the foot swings
forward.
If the hind legs were not attached to the wings they need not have
stuck out sideways. Padian t h i n k s they were held like bird legs and
that pterosaurs ran bipedally like birds (figure 8.2c). It ought to be pos-
sible to tell how the legs were held, from the structure of the hip joint,
but this is difficult because most of the fossils have the pelvis crushed.
The best fossils seem to show that the legs stuck out sideways, m u c h
as in lizards (figure 3.5a). This does not necessarily m e a n that ptero-
saurs crawled on all fours: some lizards get up on their hind legs
to run.
Most pterosaurs were small, in the size range of c o m m o n birds, but
a few were impressively large. T h e biggest k n o w n from reasonably
complete fossils is Pteranodon, which has a wing span of about 7 me-
ters (7j yds). I want you to realize h o w big this is. Try pacing out 7i
yards. (If you do it indoors you will need a big room.) The biggest wing-
span of any modern bird is only half as m u c h — 3 . 4 meters for the Wan-
dering albatross. Even so, Pteranodon was not the biggest pterosaur.
Quetzalcoatlus was even bigger w i t h a span of (probably) 12 meters,
FLYING REPTILES 109
but the fossils found so far are very incomplete. Pteranodon is the big-
gest flying animal that we know m u c h about, and m o s t of the rest of
this chapter is about it.
We cannot discuss how it lived w i t h o u t knowing something of aero-
dynamics, the basic science of flight. Figure 8.3a shows a section through
a wing of an airplane, pterosaur or bird: it does not m a t t e r which. T h e
air in front of the wing is stationary but the air behind is moving, set
in motion by the passage of the wing. Notice that this air is moving
forward and downward. It is moving forward because it has been dragged
along by the passing wing. It is moving downward largely because the
wing is tilted at an "angle of a t t a c k " to its direction of motion, but
the arched ("cambered") shape of the wing section also helps to drive
air downward. T h e air is being driven forward and downward so there
m u s t be backward and upward forces on the wing. T h e c o m p o n e n t that
acts backward along the direction of m o t i o n is called drag and the com-
ponent at right angles to the direction of m o t i o n is called lift. T h e lift
is useful (it supports the weight of the flying aircraft or animal) but
the drag is generally a nuisance. T h e lift can be made m u c h larger than
the drag by shaping the wing appropriately, and holding it at an ap-
propriate angle of attack.
Figure 8.3b shows an airplane flying horizontally. Its wings m u s t
produce enough lift to balance its weight. Its propeller blows air back-
ward, giving enough thrust to balance the drag on the wings plus the
additional drag that acts on the fuselage. Flying animals have no pro-
pellers but flap their wings in such a way as to provide thrust as well
as lift.
Figure 8.3c shows a glider. There is no propeller to give thurst but
the forces are nevertheless balanced. T h e glider is moving on a down-
ward slope, so the lift is tilted forward. T h e lift (acting upward and
forward) and the drag (upward and backward) together balance the glid-
er's weight.
Big wings can give more lift than small ones, at any particular speed.
The same wing can give more lift w h e n traveling fast, than w h e n trav-
eling slowly. T h u s lift depends on wing area and on speed. It also de-
pends on angle of attack: a bigger angle m e a n s more lift, up to a point,
but if the angle becomes too large the lift falls again. T h e m a x i m u m
F I G U R E 8.2. C o n t r a s t i n g v i e w s o f t h e s t r u c t u r e a n d p o s t u r e o f p t e r o s a u r s : (a)
Pteranodon, s h o w i n g a w i d e - w i n g e d r e c o n s t r u c t i o n o n t h e left a n d a n a r r o w -
w i n g e d o n e o n t h e r i g h t . F r o m P a d i a n 1 9 8 5 ; (b) Pteranodon r e c o n s t r u c t e d a s a
c l u m s y c r a w l e r . F r o m B r a m w e l l a n d W h i t f i e l d 1 9 7 4 ; (c) Dimorphodon r e c o n -
structed as a nimble runner, From Padian 1983. T h e length of the head was
a b o u t 1.8 m e t e r s ( i n c l u d i n g t h e c r e s t ) i n Pteranodon a n d 2 0 c e n t i m e t e r s i n Di-
morphodon.
110 FLYING REPTILES
FIGURE 8.3. (a) A vertical section through a wing, showing the forces that the
air exerts on it; (b) an airplane flying horizontally, with the forces that act on
it; (c) a glider gliding with the forces that act on it; (d) a front view of an air-
plane, showing the air that is driven downward as the wings pass through it.
lift (L ) that can be obtained by adjusting the angle of attack is pro-

max
portional to the wing area A and the square of the speed v
Lmax =
constant x Av 2
(Figure 8.4 shows h o w wing area is measured.) T h e constant depends

on the shape of the wing, both in plan and in section, but is typically
about 0.9 kilograms per cubic m e t e r for well-designed wings. (For read-
ers w h o already know about this sort of thing, this m e a n s that the
m a x i m u m lift coefficient is about 1.5.) There are some ifs and buts
here. What I have said would not be true for a very tiny wing or for a
wing moving very slowly, but is about right for bird (or pterosaur) wings,
moving in the range of speeds at which birds fly.
For an aircraft flying horizontally, the lift is fixed: it m u s t equal the
weight W of the body. This m e a n s that there is a m i n i m u m speed v„„„.
If the aircraft tries to fly slower than this, its wings cannot produce
enough lift.
FLYING REPTILES 111
W/A, the weight divided by the wing area, is called the wing loading,
so if the constant is 0.9 kilograms per cubic meter
(In this equation, speed is in meters per second and wing loading in
newtons per square meter.) This equation tells us w h y jumbo jets need
long runways. Imagine two airplanes of the same shape, one twice as
long as the other. T h e big one has eight t i m e s the volume and so, prob-
ably, eight times the weight of the small one. However, its wings have
only four times the area. Therefore it has twice the wing loading and
1.4 times the m i n i m u m speed (1.4 is the square root of two). Jumbo
jets are not the same shape as small executive jets any more than swans
are the same shape as sparrows, but the general conclusion holds: large
aircraft cannot fly as slowly as small ones and so have to taxi to higher
speeds to take off. Pteranodon was m u c h bigger than modern flying
animals. Would it have had trouble taking off?
Flying animals have an advantage over airplanes. T h e y can flap their
wings, moving t h e m rapidly through the air although the body may be
moving slowly. To take off, a small bird has only to jump into the air
and start flapping its wings, but this needs a lot of power. Large birds
cannot do this, but m u s t get their bodies moving fast before they can
take off. A large bird taking off from a cliff or branch can get up speed
F I G U R E 8.4. O u t l i n e s o f (a) a n a l b a t r o s s a n d (b) a c o n d o r , s h o w i n g t h e m e a n -

i n g s o f s o m e t e r m s . S t i p p l e i n (a) s h o w s h o w w i n g a r e a i s m e a s u r e d . A s p e c t
ratio is ( s p a n / m e a n chord).
112 FLYING REPTILES
by diving from its perch, but to take off from level ground it m u s t run
like a taxiing airplane. It helps to run into the wind because the speed
that matters is not the speed over the ground but the speed of the wings
relative to the air.
The Kori bustard, found in the East African plains, is possibly the
heaviest flying bird, with masses up to 16 kilograms. It seems to be a
big effort for it to get airborne. It has to m a k e a taxiing run, and often
simply runs away w h e n people approach, without bothering to take off.
Vultures similarly run to take off, and swans and pelicans taking off
from water run on the water surface to get up speed. All these groups
include species that reach masses of 10 kilograms or more.
This suggests that it might have been difficult for the enormous
Pteranodon to get airborne. Would it have had to run fast to take off,
and if so could it take off at all? I cannot imagine the clumsy beast
shown in figure 8.2b running, but Pteranodon m a y have moved more
like the pterosaur in Padian's reconstruction (figure 8.2c), which looks
quite fast.
It will help us to judge h o w difficult take-off would have been if we
calculate Pteranodon's m i n i m u m flying speed. For this we need to know
its wing loading, body weight divided by wing area.
We can measure the wing area from figure 8.2a, but should we use
the wide-winged reconstruction (on the left) or the narrow-winged one
(on the right)? They give very different wing areas, 4.6 and 2.5 square
meters. I will consider both possibilities.
Body weight could be calculated from the volume of a model, as has
been done for dinosaurs (chapter 2), but I do not think this has been
tried. Instead, scientists have calculated the mass from the dimensions
of bones and of drawings of the animal as they believe it looked in life,
w i t h o u t actually m a k i n g a model. T h e y got a surprizing result: Pter-
anodon'^ mass was probably only about 15 kilograms, about the same
as a Kori bustard. This is only a rough estimate and may be quite badly
wrong, but Pteranodon seems to have been a lot lighter than might
have been guessed from the huge size of its wings. Its body was small,
and remarkably lightly built.
A mass of 15 kilograms m e a n s a weight of 150 newtons. (Multiply
mass by gravitational acceleration to get weight.) This gives a wing
loading of 150/4.6 = 33 n e w t o n s per square meters for the wide wings
and 150/2.5 = 60 n e w t o n s per square meter for the narrow ones. These
give m i n i m u m speeds of meters per second for wide
wings and 8 meters per second for narrow ones.
These are remarkably low values, for such a large animal. T h e big-
gest albatrosses and vultures have smaller masses than Pteranodon but
their wing loadings are about 150 and 100 newtons per square meter,
FLYING REPTILES 1 13
respectively, giving m i n i m u m speeds of about 13 and 10 meters per

second. It may have been easier for Pteranodon to take off than it is
for these large birds. If the wind speed exceeded its m i n i m u m flying
speed it would be really easy: it would only have to face into the wind
and spread its wings, and up it would go. Winds of the necessary speed,
6 to 8 meters per second, are described by sailors as moderate breezes,
and produce small waves with fairly frequent " w h i t e horses." Alba-
trosses take off by facing the wind and spreading their wings, but they
need stronger winds because their m i n i m u m flying speeds are higher.
Pteranodon's low wing loading was partly due to the astonishing
lightness of its big wing bones. These were air-filled tubes, w i t h walls
only about a millimeter thick (figure 8.5). Tubular structure is a good
way of getting strength with lightness, w h e n bending m o m e n t s have
to be resisted, because a tube has a bigger section m o d u l u s than a solid
rod with the same a m o u n t of material in its cross section. (Section
modulus was explained in chapter 4.) This is w h y bicycle frames and
scaffolding are made of tubes. Most long bones, (including the ones in
our own bodies) are tubular, but most of t h e m are filled w i t h marrow
and so cannot be particularly light. Many bird bones are thin-walled
air-filled tubes but none have such remarkably thin walls as the wing
bones of Pteranodon. We are pretty sure that Pteranodon's bones were
air-filled because they have holes through their walls like the holes
that connect the air cavities of bird bones to the lungs.
How did Pteranodon fly once it had taken off? Most large birds spend
a lot of time gliding, using rising currents of air to keep themselves
airborne. This is called soaring, and it seems likely that Pteranodon
also soared. There are two principal soaring techniques used by differ-
ent groups of birds. They are also used by glider pilots.
F I G U R E 8.5. S e c t i o n s o f a t y p i c a l m a m m a l b o n e ( a c a m e l t i b i a ) a n d o f t h e first
p h a l a n x o f t h e l o n g f i n g e r o f Pteranodon. M o d i f i e d f r o m C u r r e y a n d A l e x a n d e r
1985.
1 14 FLYING REPTILES
O n e of these techniques is called slope soaring. When a wind blows

against a hillside or against the side of a wave it is deflected upward,
making soaring possible. Figure 8.6a shows a bird soaring along the
windy side of a wave. It is gliding, not flapping its wings, so it m u s t
be sinking relative to the air, but the air is rising. If the air is rising
fast enough the bird can travel horizontally, keeping at the same height
above the water. Albatrosses and petrels often soar along waves (but
albatross also use another soaring technique that involves swooping up
and down). Gulls similarly soar along hillsides and cliffs.
To slope soar w i t h o u t being blown downwind, a bird must glide at
least as fast as the wind. If its air speed (its speed relative to the air)
equals the wind speed, it can face directly into the wind and remain
stationary relative to the ground. To travel at right angles to the wind
it m u s t glide faster than the wind, obliquely into the wind, as shown
in figure 8.6b. (The ground speed, in this diagram, m e a n s the speed of
the bird relative to the ground.) Slope soarers have to be able to glide
fast.
T h e second important soaring technique uses thermals, which are
c o l u m n s of rising hot air. T h e sun shining on the ground heats it up,
the hot ground heats the air immediately above it and the air over the
hottest patches of ground rises as thermals. A bird gliding in a thermal
F I G U R E 8.6. D i a g r a m s of slope soaring.

FLYING REPTILES 115
gains height if its rate of sinking, relative to the air, is less t h a n the
rate at which the air is rising. T h e r m a l soaring is done by circling for
a while in a thermal, gaining height, and t h e n gliding to the next ther-
mal (figure 8.7a). Thermals are often easy to find because cumulus clouds
(the fluffy kind) form at the top of t h e m . Glider pilots find thermals
by looking for the clouds, and birds presumably do the same.
Vultures soar all day in thermals, looking for carcasses to feed on.
Some also travel long distances between nests and feeding sites by
thermal soaring. Storks migrate between Europe and Africa largely by
thermal soaring, m a k i n g large detours to avoid the Mediterranean Sea,
which has no thermals.
However, thermals do form over the sea in the parts of the tropics
and subtropics where the trade winds blow. These thermals are pro-
duced by a different m e c h a n i s m from the ones over land. T h e trade
winds blow constantly and fairly gently from the northeast in the
northern hemisphere and from the southeast in the southern hemi-
sphere, carrying cool air toward the equator. This air is heated by the
warm sea, and thermals form. Frigate birds soar in these thermals.
Thermal soaring over land or sea requires the ability to glide in small
circles, because m a n y thermals are only a few tens of meters across.
Slope soarers have to be able to glide fast and thermal soarers have
to be able to glide in small circles. We m a y get clues about Pterano-
don's flying habits by asking which of these things it could do well.
Every glider glides well only in a limited range of speeds. If it glides
slowly, near its m i n i m u m speed, it inevitably loses height rather rap-
idly. If it glides fast it again loses height rapidly. However, there is an
intermediate range of speed at which low sinking speeds (relative to
the air) are possible. I have already s h o w n that the m i n i m u m air speed
is proportional to the square root of wing loading. T h e air speed at
which the sinking speed has its lowest value is also proportional to the
square root of wing loading. This m e a n s that low wing loading is best
for slow gliding and high wing loading for fast gliding. Slope soarers,
which need to glide fast, should have high wing loading.
To glide in a circle, a bird needs a centripetal force pulling toward
the center of the circle. (Similarly, a stone whirled on the end of a
string is prevented by tension in the string from flying off at a tangent.)
Birds get the necessary centripetal force by banking (figure 8.7b), so
that the lift on their wings pulls inward (providing centripetal force)
as well as upward (balancing body weight). Plainly, the lift m u s t be
bigger than the required centripetal force which is mv /i for a glider
2
of mass m moving at speed v in a circle of radius r:
Lift is greater than mv /r.2

1 16 FLYING REPTILES
F I G U R E 8.7. D i a g r a m s of t h e r m a l soaring.
However, we already know that
M a x i m u m lift = constant x Av 1
where A is the wing area. These two s t a t e m e n t s tell us that
(constant x Av ) is greater than mv /r.

1 2
Cancel out v from both sides of this s t a t e m e n t and rearrangement it

2
to get i on the left hand side

r is greater than [(m/A) -=- constant].
This tells us that it is impossible to glide in circles of less than a
certain radius. T h e m i n i m u m possible radius is proportional to m/A
and therefore to wing loading. T h e r m a l soarers, that have to glide in
small circles, should have low wing loadings.
These arguments tell us that slope soarers should have high wing
loadings and thermal soarers should have low ones. Earlier in the chap-
ter I argued that large birds will have larger wing loadings than similar-
shaped small birds. Figure 8 . 8 a seems to show that all these things are
true. It shows that slope soarers have larger wing loadings than thermal
soarers of equal mass but that w i t h i n each group large birds have larger
wing loadings than small ones. Pteranodon has a remarkably low wing
loading for its mass, lower even t h a n for the modern thermal soarers,
whether you use the wide-winged or the narrow-winged reconstruc-
tion.
Wing loading is only one of several features of aircraft that affect
gliding performance. A n o t h e r is aspect ratio, the ratio of wing span to
m e a n chord. Figure 8.4 defines these t e r m s and contrasts the high as-
FLYING REPTILES 117
118 FLYING REPTILES
pect ratio (long, narrow) wings of an albatross with the low aspect ra-
tio (relatively short, broad) wings of a vulture. As a general rule, glid-
ers with high aspect ratios perform better than ones with lower aspect
ratios: they are capable of lower sinking speeds and of gliding at shal-
lower angles. However, it would obviously not be sensible to build wings
with a span of a few kilometers and a chord of a few millimeters: ex-
cessively large aspect ratios would mean awkwardly long wings which
would be difficult to m a k e strong enough.
I am going to explain why high aspect ratios are best, but first I will
have to explain one of the most basic laws of mechanics, N e w t o n ' s
Second Law of Motion. T h e m o m e n t u m of a moving body is its mass
multiplied by its velocity. (Please do not confuse this linear m o m e n -
t u m with the angular m o m e n t u m mentioned in chapter 5.) A force is
needed to change the m o m e n t u m of a body, and N e w t o n ' s Second Law
says that the force equals the rate of change of m o m e n t u m . Acceler-
ation is rate of change of velocity, so if mass is constant, mass times
acceleration is rate of change of m o m e n t u m . For most purposes we can
express the law in its most familiar form, force equals mass times ac-
celeration, but in this section it is more convenient to mention mo-
mentum.
T h e wings of an aircraft get lift by pushing on the air they pass
through, giving it m o m e n t u m (figure 8.3a). If they push on a mass M
of air in each unit of time, giving it downward velocity w, the mo-
m e n t u m given to the air in each unit of time is Mw. This is the rate
of change of m o m e n t u m and is equal to the lift force. T h e air is also
given kinetic energy, a m o u n t i n g to /-zMw in each unit of time. This
l 2
kinetic energy m u s t be supplied somehow, either from work done by

the engines of a powered airplane or from potential energy lost as a
glider loses height. Aircraft designers want to keep it as small as pos-
sible.
You can get the necessary lift either by pushing on a little air, giving
it a high velocity (low M, high w) or by pushing a lot of air to a low
velocity (high M, low w). The latter requires less energy. You can get
the same Mw for less /-2Mw if you m a k e M large and w small. An
l 2
airplane will need less power and a glider will lose height less fast, if
they are designed so that their wings push on as m u c h air as possible.
Figure 8.3d shows the air that the wings push on. The bigger the
wing span, the more air gets pushed. High aspect ratio (long, narrow)
wings have larger spans than low aspect ratio ones, for the same wing
area. T h a t m e a n s that, in general, high aspect ratios are best.
Figure 8.8b shows aspect ratio plotted against body mass. T h e slope
soaring albatrosses etc. have high aspect ratios and so also do the frig-
ate birds, the marine thermal soarers. However the storks, vultures,
FLYING REPTILES 1 19
etc., that soar in thermals over land, have relatively low aspect ratios.
One possible reason is that longer, higher aspect ratio wings, that would
give better gliding performance, might be awkward when the bird was
taking off from land. T h e point for the wide-winged reconstruction of
Pteranodon, in figure 8.8b, is between the lines for slope soarers and
land thermal soarers, but the point for the narrow-winged one is above
the line for slope soarers.
Whether the wings of Pteranodon were broad or narrow, the wing
loading is low enough to suggest that it soared in thermals. It is not
so clear whether it soared inland or over the sea. T h e wide-winged re-
construction is not too different from the large land soarers, with an
aspect ratio only a little higher than theirs and w i t h an even lower
wing loading than any of t h e m have. T h e narrow-winged model seems
like a larger version of the frigate birds, marine soarers w i t h high as-
pect ratios and low wing loadings. I have already said that I prefer the
narrow-winged reconstruction, on anatomical grounds. N o w we will
test the hypothesis, that Pteranodon was a marine thermal soarer.
The hypothesis suggests that Pteranodon fossils should be found in
rocks formed from sediments laid down in the sea. T h e y are actually
found in the central and western United States and southern Russia,
mostly in places that are now well away from the sea. However, the
same rocks contain fossils of sharks and other fishes, plesiosaurs (see
Chapter 9) and turtles so it seems clear that they were formed in the
sea and that coastlines have been moved around by subsequent earth
movements.
This is not enough to show that the hypothesis is plausible. It is
only in the trade wind zones that there are likely to be enough ther-
mals over the sea for soaring to be feasible. Nowadays the trade winds
blow only in and near the tropics, from about 25°N (the latitude of
Miami) to 25°S (a little south of Rio de Janiero). Pteranodon is found
further north. It is particularly abundant in Kansas (about 40°N) but is
also found in Alberta (55°N). T h e c o n t i n e n t s have moved a little over
the earth's surface since the Cretaceous period, but the latitudes of the
Pteranodon sites have not been changed m u c h . There is a lot of evi-
dence that climates were generally w a r m e r than they are now, but I
have not been able to find any reliable opinion as to how far north it
would have been possible to soar in thermals over the sea.
The hypothesis, that Pteranodon was a marine thermal soarer, also
suggests that it would have eaten food from the sea. Fish scales and
bones have been found in some fossils, in the position of the stomach.
Another fossil had remains of fish and a leg of a crustacean in its throat.
Pteranodon probably fed mainly on fish, and its bird-like beak seems
suitable for catching them, though perhaps rather narrow.
120 FLYING REPTILES
Pteranodon seems m u c h too fragile to have dived into the water to

catch fish. It seems m u c h more likely that it fed like frigate birds, flying
low over the water and grabbing fish or squid without landing, putting
only its beak into the water. T h e narrowness of the beak might ac-
tually be an advantage, because it would reduce the drag of the water
on it. Pteranodon m a y have kept itself airborne by thermal soaring,
searching for prey s w i m m i n g close to the. surface of the sea and swoop-
ing down to catch t h e m .
T h e evidence does not prove that Pteranodon flew and fed like a
giant frigate bird, but it seems consistent w i t h the idea. Thermals over
the sea continue day and night, in the trade wind zone, so Pteranodon
could have soared for days on end w i t h o u t landing, seldom having to
flap its wings. It could not have flown very fast because of its low wing
loading, but if a storm blew up it could have avoided being blown
downwind by landing on the surface of the sea.
T h e long beak may have been good for catching fish but seems likely
to have been troublesome in flight. All would be well so long as it faced
directly forward, but if it was turned to either side the air would press
on one side of the beak and try to twist the head round like a weather
cock. Pteranodon would need strong neck muscles, unless it had some
way of canceling the effect out.
Cherrie Bramwell and G. R. Whitfield of the University of Reading
thought that the crest on the back of Pteranodon's head (figure 8.2b)
might have helped to cancel out the twisting effect, keeping the head
facing forward. They tested the idea by m a k i n g models of the head,
with and w i t h o u t the crest, and putting t h e m in a wind tunnel. They
set t h e m at various angles to the wind and measured the m o m e n t s that
the aerodynamic forces exerted on t h e m . They found to their surprize
that the crest had little effect.
Different species of Pteranodon have the crest set at different angles
and most pterosaurs (including some large ones w i t h long beaks) have
no crest. It seems possible that the m a i n function of the crest had noth-
ing to do w i t h aerodynamics. It may have been an ornament with the
same sort of function as the crests of hadrosaurs (chapter 5). It has also
been suggested that it m a y have been a cooling fin like the plates on
the backs of stegosaurs (chapter 7).
This chapter has concentrated on Pteranodon, the largest flying an-
imal k n o w n from reasonably complete fossils. It was enormous, with
a wing span of 7 meters, but very lightly built. We know the span of
its wings, but paleontologists disagree about their shape (figure 8.2a).
Whether the wings were wide or narrow, its wing loading was appar-
ently low, which would have enabled it to fly slowly and suggests that
it soared in thermals. T h e wide wings of one restoration resemble the
FLYING REPTILES 121
wings of vultures, which soar in thermals over land. T h e narrow wings

of the other are like those of frigate birds, which soar in thermals over
the sea in the trade wind zone. It seems likely that Pteranodon flew
like frigate birds and fed like them, grabbing fish from the surface of
the sea.
Principal Sources
B r a m w e l l a n d W h i t f i e l d (1974) i n v e s t i g a t e d t h e a e r o d y n a m i c s o f t h e w i d e - w i n g e d
r e s t o r a t i o n of Pteranodon a n d B r o w e r (1983) did t h e s a m e for t h e n a r r o w - w i n g e d
r e s t o r a t i o n . P a d i a n (1983, 1985) a r g u e d t h a t t h e n a r r o w - w i n g e d r e s t o r a t i o n w a s t h e
m o r e r e a l i s t i c . P e n n y c u i c k (1983) i n v e s t i g a t e d t h e flight of frigate birds a n d R a y n c r
(1987) h a s s h o w n h o w t h e w i n g l o a d i n g s a n d a s p e c t r a t i o s o f b i r d s are r e l a t e d t o
t h e i r s t y l e s of flying. W c l l n h o f e r (1975) is t h e s o u r c e for figure 8 . 1 .
B r a m w e l l , C. D. and G. R. W h i t f i e l d . 1974. B i o m e c h a n i c s of Pteranodon. Philo-

sophical Transactions of the Royal Society B 2 6 7 : 5 0 3 - 5 8 1 .
Brower, ]. C. 1983. T h e a e r o d y n a m i c s of Pteranodon a n d Nyctosaurus, t w o large
p t e r o s a u r s from t h e U p p e r C r e t a c e o u s of K a n s a s , journal of Vertebrate Palaeon-
tology 3 : 8 4 - 1 2 4 .
C u r r e y , J. D. a n d R. Mc N. A l e x a n d e r . 1985. T h e t h i c k n e s s of t h e w a l l s of t u b u l a r
b o n e s , lournal of Zoology A 2 0 6 : 4 5 3 - 4 6 8 .
Padian, K. 1983. A f u n c t i o n a l a n a l y s i s of flying a n d w a l k i n g in p t e r o s a u r s . Paleo-
biology 9 : 2 1 8 - 2 3 9 .
Padian, K. 1985. T h e o r i g i n s a n d a e r o d y n a m i c s of flight in e x t i n c t v e r t e b r a t e s . Pa-
laeontology 28:413-433.
P e n n y c u i c k , C . I . 1983. T h e r m a l s o a r i n g c o m p a r e d i n t h r e e d i s s i m i l a r t r o p i c a l bird
species, Fregata magnificens, Pelecanus occidentalis a n d Coragyps atralus. jour-
nal of Experimental Biology 1 0 2 : 3 0 7 - 3 2 5 .
R a y n e r , J. M. V. 1987. F o r m a n d f u n c t i o n in a v i a n flight. Current Ornithology 5:
1-66.
Wcllnhofer, P . 1975. D i e R h a m p h o r h y n c h o i d e a der O b e r i u r a P l a t t e n k a l k e Sud-
d e u t s c h l a n d s . Palaeonlographica A 1 4 8 : 1 - 3 3 , 1 3 2 - 1 8 6 , a n d 1 4 9 : 1 - 3 0 .
IX
Marine Reptiles
W E WILL now discuss the giant reptiles that lived in the sea in the
Mesozoic era, w h e n the dinosaurs were living on land. They had
flippers instead of feet and would probably have been pretty helpless
on land. We k n o w that they lived in the sea rather than in fresh water
because the other fossils found in the same rocks include sea urchins
and squid-like molluscs, m e m b e r s of groups whose modern members
are found only in the sea.
I am going to start w i t h the ichthyosaurs—reptiles that looked re-
markably like fish (figure 9.1). Most of t h e m were at least a meter long
and some were as m u c h as 15 meters. I have a model of one of the
larger kinds, Ichthyosaurus, bought from the Natural History Mu-
seum, London. I measured its volume and calculated the mass of the
living animal in the same way as for dinosaurs (chapter 2). This par-
ticular animal was 8 meters long and I calculate that its mass was 6
tonnes. Adult Killer whales have about the same length and mass.
Most ichthyosaur fossils are skeletons and nothing more, but a few
have dark m a r k s showing the outline of the body and of the fins and
flippers. These m a r k s show that the flippers were a good deal broader
than you might guess from the skeletons, and that at least some ich-
thyosaurs had a fin on the back. Some ichthyosaurs had straight ta-
pering tails but the best k n o w n kinds, including the one in figure 9.1,
had a sharp kink in the backbone where it entered the tail fin. Dark
m a r k s on a few fossils show that these ichthyosaurs had tails shaped
like crescent m o o n s .
There are t w o groups of modern fish that look very like these ich-
thyosaurs: the t u n n i e s and the porbeagle sharks (figure 9.2). As well as
being shaped like ichthyosaurs they overlap the ichthyosaur size range.
For example, Bluefin tuna reach lengths of 4 meters and masses of 0.8
tonnes. Great w h i t e shark grow to m a x i m u m lengths of about 11 me-
MARINE REPTILES 123
FIGURE 9.1. T h e skeleton of a Jurassic ichthyosaur, w i t h an o u t l i n e of the

body. F r o m R o m e r 1966. I h a v e added a frogman to scale, a s s u m i n g t h i s ich-
t h y o s a u r is 8 m e t e r s long.
ters. Whales also look like ichthyosaurs, but w i t h a very obvious dif-
ference. Ichthyosaurs had vertical tail fins (like t u n n i e s and sharks and
presumably beat t h e m from side to side w h e n they swam. Whales have
horizontal tail flukes and beat t h e m up and down.
In one respect ichthyosaurs were even more like dolphins than like
tunnies or sharks: they had long narrow jaws with a lot of simple pointed
teeth. Dolphins eat fishes and squid, and ichthyosaurs seem to have
eaten similar things. Many ichthyosaurs have been found w i t h their
fossilized stomach contents still in place inside t h e m , enclosed by their
ribs. Some fish scales have been found in them, and e n o r m o u s n u m b e r s
of hooks from the suckers of squid-like m o l l u s k s .
Tunnies, porbeagle sharks, and whales, the modern animals shaped
like ichthyosaurs, swim fast. T h e m o s t reliable speed m e a s u r e m e n t s
have been made with dolphins trained to s w i m as fast as possible over
a marked course, or to follow a lure towed by a fast boat. T h e highest
speed on record seems to be 11 meters per second (25 mph) for a Spot-
ted porpoise 2 meters long, and slightly slower speeds have been re-
corded for other species. These are sprint speeds, maintained for only
a few seconds. They are astonishingly fast for m o v e m e n t in water, and
equal the top speeds (on land) of h u m a n sprinters.
Speeds of tunnies have been measured, both by filming t h e m and by
catching t h e m on a rod with an i n s t r u m e n t e d reel, that recorded the
rate at which the fish pulled out the line. Several records of 5 to 13
meters per second were obtained in this way, and also two of 21 meters
per second for a Wahoo and a Yellowfin tuna. I find these last t w o
records hard to believe because they are so m u c h higher than any oth-
ers. I wonder whether some error was made: for example, a mistake
could conceivably have been made about the speed at which the re-
124 MARINE REPTILES
F I G U R E 9.2. A Porbeagle s h a r k , a Bluefin t u n a , a n d an o p t i m u m s t r e a m l i n e d

s h a p e . T h e fishes are from d r a w i n g s by Valerie du H e a u m e in W h e e l e r 1969.
cording e q u i p m e n t was running. Even if we reject these records (as I

am inclined to do) it seems clear that tunas, like dolphins, swim very
fast. T h e fastest shark speed I have seen recorded is 5 meters per sec-
ond, but there are very few data.
The speeds of the similar-shaped modern animals make it seem likely
that ichthyosaurs were also fast. My guess is that they could have
sprinted at 10 meters per second.
Drag resists the m o v e m e n t of bodies through water, as also through
air. It is m u c h larger in water t h a n at the same speed in air because
water is so m u c h denser. To m i n i m i z e drag, a body should be designed
to disturb the water as little as possible, as it passes through. Any body
will leave a wake of swirling water behind it, but the narrower the
wake, the less the drag. This is because energy is needed to set the
water swirling: the kinetic energy of the swirling water comes from
work done against drag. Streamlining is the art of designing bodies so
as to disturb the water as little as possible. The best shapes are rounded
in front, and taper to a fine point behind to allow the water to close
MARINE REPTILES I 2^
in smoothly after the body has passed. Torpedoes and submarines are
shaped like this, and so also are ichthyosaurs, whales, and m a n y fish.
When airships were used, the engineers w h o designed t h e m set out
to discover the best shape. The carrying capacity of an airship depends
on its volume because it is supported by the buoyancy of gases that
are lighter than air, so the basic problem was to find the shape that
gave least drag for given volume, at any particular speed. T h e answer
turned out to be a streamlined shape with the length 4.5 t i m e s the
diameter at the fattest part (figure 9.2, bottom).
The same shape seems likely to be best for s w i m m i n g animals, so
it is not surprizing to find that ichthyosaurs, tunnies etc. are very nearly
this shape. The Ichthyosaurus model (already mentioned) has its length
5.0 times the m a x i m u m diameter, Yellowfin tuna are 4.5 diameters
long, and Porbeagle sharks and Bottle-nosed dolphins are both about
5.5 diameters long. (The diameter that I have used in these calculations
is the mean of the m a x i m u m height of the body and the m a x i m u m
width.)
Figure 9.3 shows how tunnies swim. T h e y beat their tails from side
to side as they move forward, so the tail takes a wavy path through
the water. It is held at an angle of attack so that lift acts on it as well
as drag. (Lift acts on hydrofoils in water, just as on aerofoils in air.)
While the tail is moving to the right, the lift acts forward and to the
left. While it is moving to the left, the lift acts forward and to the right.
The components to left and right cancel out, over a complete cycle of
tail movements, so the net effect is a forward thrust, driving the fish
through the water. T h e drag on the tail acts backward all the time,
reducing the thrust, but if the hydrofoil is well designed (as t u n n y tails
seem to be) the drag is relatively small. Ichthyosaurs like the one in
figure 9.1 presumably swam like this but it has been suggested that
some of the ones w i t h narrow tapering tail fins m a y have depended
more on flipper m o v e m e n t s .
FIGURE 9.3. How tunnies swim. Ichthyosaurs presumably s w a m in the same

way.
126 MARINE REPTILES
T h e aspect ratios of tails can be measured in the same way as for

wings, by dividing the span of the tail by its mean chord (figure 8.4).
High aspect ratio tails give the same lift for less drag, like high aspect
ratio wings, and m a k e for efficient s w i m m i n g . T u n n y tails have high
aspect ratios, for example 7 for Yellowfin. Whale flukes have lower
aspect ratios, for example 5 for White-sided dolphin, possibly because
they have no skeleton to stiffen t h e m and might be too flexible if their
spans were increased. Tail outlines preserved in ichthyosaur fossil show
still lower aspect ratios, for example 3.7 for Ichthyosaurus. In this re-
spect ichthyosaurs seem markedly inferior to tunnies.
As well as being similar in shape, tunnies, porbeagle sharks, and
whales have a remarkable thing in c o m m o n . All of t h e m are endo-
t h e r m s . You would expect whales to be endotherms, because they are
m a m m a l s , but t u n n i e s and porbeagle sharks seem to be the only en-
dothermic fishes. They do not keep themselves as w a r m as m a m m a l s ,
and they do not heat the whole body to the same temperature: the
warmest parts deep inside the body are seldom above 35°C and often
m u c h cooler, whereas m a m m a l s keep their bodies at 3 6 - 4 0 ° C . How-
ever, these w a r m e s t parts are often 10 to 15 degrees warmer than the
water in Bluefin tuna, and 10 degrees in Porbeagle sharks. It seems
quite likely that ichthyosaurs were also endotherms.
Dolphins often leap repeatedly from the water, as they swim: this
is called "porpoising." It looks as if they are playing or exercising, but
it has been suggested that they m a y actually save energy by this ap-
parently strenuous behavior. T h e m a i n point is that drag is very m u c h
less in air than in water, so if the dolphins can m a k e part of their jour-
ney out of the water they may save energy.
I will try to explain the idea in a bit more detail, because it seems
possible that ichthyosaurs also porpoised. Obviously, it costs energy to
leap: the a m o u n t of energy is proportional to the height of the leap.
T h e length of the leap depends on the dolphin's speed and on the angle
at which it leaves the water, but faster take-off (at any particular angle)
makes the leap longer and also higher. If leaps at the same angle are
compared, the height of the leap (and so the energy needed) is propor-
tional to the length of the leap, so the energy needed per meter leaped
is the same for all s w i m m i n g speeds.
A leap saves energy if the work needed for it is less than would be
needed to swim the same distance. T h e work needed for s w i m m i n g is
done against drag, and can be calculated by multiplying the drag by the
distance. (The work done against a force is the force multiplied by the
distance moved against it.) Drag is roughly proportional to the square
of speed, so the work needed to swim a meter increases w i t h increasing
speed, while the work needed to leap a m e t e r remains constant. As a
MARINE REPTILES 127
dolphin swims faster and faster it m u s t eventually reach a speed at

which leaping saves energy.
People have tried to calculate the critical speed, and one estimate
for dolphins is 5 meters per second, which is well w i t h i n their range
of speeds. However, there are all sorts of doubts about the calculation,
both about the amount of work needed for leaping and about the amount
needed for s w i m m i n g . All we can feel sure of is that there m u s t be
some speed, above which leaping would save energy if dolphins can
swim that fast. If dolphins leap for this reason, perhaps ichthyosaurs
leapt too. I like to imagine that they did, but tunnies give no support
to the idea. They do not leap although they are m u c h the same size
and shape as dolphins and seem able to swim as fast.
Most bony fish have a gas-filled float (the swimbladder) in the body
cavity, making their densities about the same as the density of the
water they swim in. Sharks and most tunnies have no swimbladder,
so they are denser than water and would sink if they stopped swim-
ming. Porbeagle sharks and tunnies live near the surface of the oceans,
often with the b o t t o m thousands of m e t e r s below, so there is no ques-
tion of stopping for a quick rest on the bottom. These fish swim all
the time, and some apparently have to keep s w i m m i n g in any case, to
get the oxygen they need. (Instead of m a k i n g breathing m o v e m e n t s like
other fish they simply swim around with their m o u t h s open, letting
the water flow through their gills.) They prevent themselves from sink-
ing mainly by keeping their pectoral fins spread like airplane wings, at
a suitable angle of attack to give the necessary lift. (The pectoral fins
are the large pair close behind the head.)
Whales are less dense, partly because they have a lot of blubber (fat
is slightly less dense than water) but largely because they have air-
filled lungs. Whales can often be seen floating at the surface of the sea
with their backs protruding slightly above the surface, showing that
they are actually less dense than the water. Their buoyancy enables
t h e m to stop s w i m m i n g and rest w i t h o u t sinking. Ichthyosaurs pre-
sumably also had lungs (it would be surprising to find a reptile w i t h o u t
them) and probably also had densities close to the density of water.
Similarly crocodiles have about the same density as (fresh) water, as I
showed in chapter 2. Ichthyosaurs probably had no need to keep swim-
ming like tunnies and Porbeagle sharks, but could stop and rest like
whales.
Ichthyosaurs m u s t have come to the surface regularly to breathe, but
they may also have dived to substantial depths, like whales. If they
had the same density as water while at the surface, they would be den-
ser while diving, because the air in their lungs would be compressed
to a smaller volume. T h e pressure at a depth of 10 meters is twice the
128 MARINE REPTILES
pressure at the surface and would halve the volume of the air at 20 ;
meters the pressure is three t i m e s as m u c h as at the surface and the

volume of the air would be reduced to one third; and so on. During a
deep dive the lungs would give very little buoyancy and the animal
would sink if it stopped s w i m m i n g . It might get the necessary lift, as
it swam, by spreading its flippers.
Ichthyosaurs may have had to dive quite deep, to get their food. Their
skulls show that they had large eyes, which suggests that they de-
pended on sight to find prey. Further, it suggests to me that they fed
by day (but I admit that big eyes could be an adaptation for feeding in
dim light, at dusk). Many fish and squids spend the night near the sur-
face but dive quite deep by day: for example, herring that spend the
night near the surface often dive to 100 meters or more. If ichthyosaurs
fed by day on prey that behaved like that, they would have had to dive.
T h e ichthyosaurs seem splendidly adapted for swimming, but they
would probably have been as helpless on land as a stranded whale. They
could hardly have crawled up beaches to lay eggs above the high tide
mark, as the sea turtles do. They could not have laid their eggs in water,
because the embryos would have suffocated, as the embryos of birds
and modern reptiles do if their eggs are submerged. T h e reason is that
oxygen diffuses m u c h more slowly through water than through air. If
the pores of the eggshell get filled w i t h water, oxygen cannot diffuse
in fast enough.
Ichthyosaurs seem to have got round this problem by giving birth
instead of laying eggs. Adults have been found with up to 12 young
ones fossilized inside them, enclosed by their ribs. It is possible that
they had eaten the little ones, but it is pleasanter to think that they
were pregnant. Though most modern reptiles lay eggs, some sea snakes
and others give birth.
T h e mosasaurs were another group of m a r i n e reptiles, but they looked

m u c h less fish-like than the ichthyosaurs, more like crocodiles with
flippers instead of legs. T h e details of their skulls show that they were
actually lizards, closely related to the Komodo dragon and other mon-
itor lizards. T h e biggest of t h e m were at least 9 meters long, about the
same as the biggest modern crocodiles. (Though the Komodo dragon is
the largest modern lizard it grows little more than 3 meters long.) T h e
mosasaurs lived in the Cretaceous period, at the end of the Mesozo-
ic era.
I speculated that ichthyosaurs m a y have dived, and there is some
evidence that mosasaurs did. X-ray pictures of many of their vertebrae
show the kind of damage that would have occurred if the blood supply
to parts of the bone had got cut off, while the animal was still alive.
MARINE REPTILES 129
It has been suggested that the damage m a y have been caused by the
bends, a serious hazard of diving.
Here is how the bends happens to h u m a n divers. The air they breathe
has to be compressed to m a t c h the pressure of the water where they
are working. T h e extra pressure m a k e s extra gas dissolve in the blood.
When the diver returns to the surface the extra gas comes out of so-
lution, forming bubbles that may block blood vessels, causing damage,
pain and even death. H u m a n divers avoid the bends by returning slowly
to the surface but a diving mosasaur would have to get to the surface
reasonably soon, for its next breath. Whales avoid the bends largely by
having small lungs and a big windpipe: the high pressures that they
meet when they dive collapse their lungs, forcing the air back into the
windpipe whether there is less danger of too m u c h gas being absorbed
into the blood.
T h e plesiosaurs lived in the Mesozoic seas, like ichthyosaurs and m o -

sasaurs. Also like ichthyosaurs and mosasaurs they had two pairs of
flippers instead of the fore and hind legs of most other reptiles. T h e
shapes of their bodies, however, were quite different from those of the
other marine reptiles. T h e t r u n k was broad and relatively low, a less
extreme version of the body shape seen in modern turtles. Attached to
it was either a long neck with a small head, or a short neck with a
relatively large head (figures 9.4, 9.5). Some of t h e m were very large,
as the picture shows, but even Elasmosauius was only half as long as
F I G U R E 9.4. A s h o r t - n e c k e d p l e s i o s a u r [Kronosaurus] a n d a l o n g - n e c k e d o n e
(Elasmosaurus) f r o m R o m e r 1 9 6 8 , w i t h a f r o g m a n .
130 MARINE REPTILES
F I G U R E 9 . 5 . S k e l e t o n s o f t w o l o n g - n e c k e d p l e s i o s a u r s . Cryptoclidus ( l e n g t h 4
m e t e r s ) i s s h o w n i n s i d e v i e w a n d Thaumatosaurus ( l e n g t h 3.4 m e t e r s ) i s s h o w n
from below. From Brown 1981, by courtesy of the British M u s e u m (Natural
History), a n d R o m e r 1966, respectively.
the dinosaur Diplodocus. T h e Natural History M u s e u m , London, sells

models of a very large (14 meter) long-necked plesiosaur. O n e of my
students, Debbie O'Hare, measured its volume and calculated that the
living animal had a mass of 7.5 tonnes (or perhaps a little less: the
model is possibly a little too deep in the body). In comparison with
that, the Leatherback, the biggest of the modern sea turtles, grows to
shell lengths of only 2 meters and masses of about 0.6 tonnes.
Plesiosaurs seem to have had small tails that would not be very ef-
fective for s w i m m i n g . They presumably propelled themselves mainly
by flipper m o v e m e n t s , either by rowing (like freshwater turtles) or by
"underwater flight" (like marine ones).
Figures 9.6a and 9.7a show how plesiosaurs may have rowed. T h e y
show the flippers moving backward and forward, like the oars of a boat.
In the backward stroke the blades of the flippers are held vertical, so
as to push as hard as possible against the water. In the forward stroke
they are held horizontal so as to strike the water edge-on and meet as
little resistance as possible. (A h u m a n rower would lift the oars out of
the water for the forward stroke, but an animal s w i m m i n g below the
MARINE REPTILES 131
surface cannot do that.) Notice that in the power stroke the flipper
blade is moving backward through the water so the drag on it (the force
opposite to the direction of m o v e m e n t ) acts forward. Rowing boats and
rowing animals are propelled by forward-acting drag on backward-mov-
ing oars.
Figures 9.6b and 9.7b show a different m e t h o d of swimming, which
is called underwater flight because the m o v e m e n t s are like those of
flying birds. Penguins, which cannot fly in air, use their wings in this
way to swim underwater. Sea turtles also swim this way, using their
flippers. T h e flippers beat up and down. On the downstroke they are
held at an angle of attack so that lift acts on them, forward and upward
(figure 9.7b). For the upstroke the angle is adjusted so that the lift acts
forward and downward. T h e upward and downward c o m p o n e n t s cancel
out over the complete cycle of m o v e m e n t s so that the net effect is a
forward thrust (which is reduced a bit by the drag on the flipper). No-
tice how similar figure 9.7b is to the diagram of a t u n n y s w i m m i n g in
figure 9.3. T u n n i e s and presumably ichthyosaurs swim by m e a n s of
their tails, beating t h e m from side to side, and turtles and possibly
plesiosaurs swim by means of their flippers, beating t h e m up and down,
but the basic principle is the same in both cases.
I would like to emphasize that rowing and underwater flight are ut-
terly different techniques. In rowing, the flippers or oars are moved
backward and forward and the propulsive thrust comes from the drag
on t h e m in their backward strokes. In underwater flight the m o v e m e n t
is up and down and the thrust comes from lift: in this case, the drag
is simply a hindrance.
I have been assuming that plesiosaurs had about the same density
as water, so that an upward force at one stage of the cycle of flipper
m o v e m e n t s m u s t be balanced by a downward force at another. In figure
9.7b this balance comes from an upward-sloping force in the down-
stroke and a downward-sloping force in the upstroke. Figure 9.7c shows
another possible way of avoiding unbalanced vertical forces. T h e flip-
per is moved almost vertically downward, held at an angle of attack
so as to give forward lift. It is then raised on a sloping path, moving
edge-on, with no angle of attack, so that the forces on it are very small.
The downstroke gives horizontal thrust and the upstroke very little
force. Figure 9.6c shows on the left how the flippers would have to
move relative to the animal's trunk, to follow the appropriate path
through the water: they would have to beat down and back, t h e n for-
ward and up. T h e thrust, in this s w i m m i n g technique, comes from lift,
so it is a form of underwater flight. Sea lions seem to swim rather
like this.
Plesiosaurs would have had to drive water backward, to propel them-
132 MARINE REPTILES
F I G U R E 9 . 6 . T h r e e p o s s i b l e s w i m m i n g t e c h n i q u e s for p l e s i o s a u r s : (a) r o w i n g ;
a n d (b) a n d (c) u n d e r w a t e r f l i g h t . T h e d i a g r a m s o n t h e left s h o w h o w t h e flip-
pers w o u l d have been m o v e d relative to the body and those on the right s h o w
s u c c e s s i v e p o s i t i o n s o f t h e a n i m a l m o v i n g t h r o u g h t h e w a t e r . O n l y t h e fore
flippers are s h o w n .
selves forward. In rowing, the flippers would push fairly small lumps
of water backward (figure 9.8a). In underwater flight the flippers, beat-
ing up and down through a large angle, would affect m u c h more water
(figure 9.8b). When I wrote about the aspect ratios of wings (chapter 8)
I explained that less energy is needed to get a force by accelerating a
large m a s s of fluid to a low velocity, than by accelerating a small mass
to a high one. T h i s argument says that underwater flight should be
more economical than rowing. It should need less energy for swim-
ming at the same speed.
T h a t is one reason for thinking underwater flight more likely than
rowing. Animals tend to evolve efficient ways of doing things. There
are animals that row, but at least some of t h e m (freshwater turtles,
and ducks) use their legs for walking as well as for rowing. It would
MARINE REPTILES 133
F I G U R E 9.7. D i a g r a m s s h o w i n g a s e c t i o n t h r o u g h a flipper, a n d t h e forces act-

i n g o n it, a t d i f f e r e n t s t a g e s o f t h e t h r e e s w i m m i n g t e c h n i q u e s o f f i g u r e 9 . 6 .
be very difficult to design a foot which was both effective for walking
on land and suitably streamlined for use in underwater flight.
Another reason for thinking underwater flight more likely is that
plesiosaur flippers taper at the tips. There is no advantage in tapering
the tip of an oar blade, and the oars used in rowing races are made
with square ends. However, there is an advantage in giving aerofoils
and hydrofoils tapered, rounded ends: such shaping can spread the lift
out over the span of the hydrofoil in the best possible way, so as to
get lift with as little drag as possible. Bird and airplane wings, and pro-
peller blades, generally taper toward their tips. T h e shape of plesiosaur
flippers suggests that their function was to provide lift, not drag.
For underwater flight, plesiosaurs would have to have been able to
flap their flippers up and down. T h e shapes of the joints seem to show
that they could have done this, though they probably could not have
raised the flippers very high above their backs. They would also have
134 MARINE REPTILES
FIGURE 9 . 8 . A plesiosaur (a) rowing and (b) "flying" under water. Broken out-
lines show the water driven backward by the swimming movements.
needed appropriate muscles. Scientists have tried to work out how the
flipper muscles were arranged, by looking at plesiosaur skeletons. Fig-
ure 9.5 shows big plates of bone to which muscles could have attached
in the chest (between the fore flippers) and on the underside of the
abdomen (between the hind flippers). These are the ventral parts of the
pectoral and pelvic girdles. T h e y seem excellent areas of a t t a c h m e n t
for muscles that would pull the flippers down, in a powerful down-
stroke, but there does not seem to be m u c h to attach upstroke muscles
to. T h e big plates of bone are all below the shoulder and hip joints.
T h e upward extensions of the pectoral and pelvic girdles (seen in the
side view) seem to be attached rather weakly to the ribs and backbone.
Strong upstroke muscles could have been attached to the backbone,
but there is little to prevent t h e m pulling the girdles bodily upward
instead of flapping the flippers. T h e symmetrical style of underwater
flight, shown in figures 9.6b and 9.7b, needs equally strong upstroke
and downstroke muscles. T h e style shown in figure 9.6c and 9.7c needs
big forces for the downstroke but only small forces for the upstroke,
and seems the more likely s w i m m i n g technique for plesiosaurs.
Plesiosaurs probably could not s w i m very fast. One reason for think-
ing this is that the v o l u m e of flipper muscle that it seems possible to
fit into their bodies is relatively small, compared to the volume of tail
MARINE REPTILES 135
muscle that would be found in similar-sized fishes or whales. T h e

swimming muscles of a 43-kilogram porpoise had a mass of 9 kilo-
grams. (This includes the tail m u s c l e and most, but not all, of the back
muscle.) That is 21 percent of body mass. I do not see how the flipper
muscles of plesiosaurs could have been as m u c h as 21 percent of
body mass.
Another reason for thinking that plesiosaurs were probably rather
slow is that the s w i m m i n g technique we suspect they used (figure 9.7c)
requires the flipper to be moved almost vertically down through the
water. To do this while the animal was moving forward, the flippers
would have to move backward (relative to the body) as fast as the body
was moving forward (relative to the water). Imagine the plesiosaur shown
in figure 9.7c s w i m m i n g at 10 meters per second, about the m a x i m u m
sprinting speed of tunnies and dolphins. During the power stroke, a
point at the center of the flipper would have to move backward through
a distance x at about 10 meters per second. If the animal was a mod-
erate-sized short-necked plesiosaur, three meters long, x would be a
meter or less and the m o v e m e n t would have to be made in one-tenth
of a second. If the upstroke was made at the same speed, the cycle of
flipper m o v e m e n t s would be completed in 0.2 seconds and the flapping
frequency would be 1/0.2 = 5 cycles per second. I doubt w h e t h e r so
large an animal could have managed so high a frequency of m o v e m e n t .
Small penguins beat their wings w h e n they swim at up to 4 cycles per
second, but they are very m u c h smaller. There is a general rule that
large animals cannot move their limbs at as high frequencies as small
ones: a horse cannot m a k e as m a n y strides per second as a mouse, and
a swan cannot m a k e as m a n y wingbeats per second as a sparrow. Figure
9.9 shows some data. Penguin wing beat frequencies are about the same
as the wing beat frequencies of similar-sized flying birds (although pen-
guins move their wings in water) but considerably less t h a n the stride
frequencies of similar-sized m a m m a l s , which is not surprizing: the
penguins were moving their wings in water. T h e points are widely
scattered around the lines but the general trend is clear: bigger animals
use lower frequencies. By extending the penguin line, I estimate that
an underwater flier 3 meters long (the size of the plesiosaur I have been
discussing) would have beaten its flippers at a m a x i m u m frequency of
about 1 cycle per second. This is only one fifth as m u c h as I estimated
would be needed for s w i m m i n g at 10 meters per second and suggests
that the plesiosaur could only manage about 2 meters per second.
It is not just chance that m a k e s big animals move their limbs at

lower frequencies than small ones: there is a good mechanical reason.
Imagine two animals of the same shape, one twice as long as the other
(and twice as wide and twice as high). It is eight t i m e s as heavy as the
136 MARINE REPTILES
small one and has eight times as m u c h limb muscle, able to do eight
times as m u c h work to accelerate the limbs at the beginning of each
stroke. This work is used to give the limbs kinetic energy (half mass
t i m e s speed squared) but its limbs, plus any water moved by them,
have eight t i m e s the mass of the smaller animal's ones, so can only
be accelerated to the same speed. The big animal's limbs have to travel
twice as far as the small ones to m a k e each stroke, so their cycle of
m o v e m e n t s takes twice as long. This argument says that doubling the
length of an animal should halve its frequency of limb movements, and
figure 9.9 shows that this is roughly true for flying birds and s w i m m i n g
penguins.
I have argued that the style of underwater flight shown in figure 9.6c
is unlikely to be very fast, because the flippers have to move back,
relative to the body, as fast as the body advances through the water.
Penguins and sea turtles use more symmetrical styles, like the one
shown in figure 9.6b which is not limited in this way, but even so they
MARINE REPTILES 13 7
are not very fast. The highest speed s h o w n in films of penguins swim-
ming in Detroit Zoo was 3.4 m e t e r s per second (for a King penguin,
about 90 centimeters long) and adult Green turtles seem to swim no
faster than 2.0 meters per second.
Penguins and sea turtles move their left and right wings or fore flip-
pers in unison. If they did not they would waste energy by s w i m m i n g
a slightly zigzag route. Plesiosaurs probably also moved the left and
right flippers of a pair together. Sea turtles swim mainly with their big
fore flippers, beating their small hind flippers only occasionally. Ple-
siosaurs had big hind flippers as well as big big fore ones and probably
used both pairs about equally.
I have argued that the downstroke was the power stroke. If so, there
might be an advantage in beating the fore and hind flippers out of phase
with each other as shown in figure 9.8b. Fore-power strokes would al-
ternate with hind-power strokes, keeping the animal moving at a steady
speed. This might save energy; drag is about proportional to speed
squared, so the average drag is greater w h e n s w i m m i n g at a fluctuating
speed than when s w i m m i n g steadily at the same average speed. How-
ever, there might be a very serious disadvantage in beating the fore and
hind flippers alternately. T h e hind flippers might find themselves mov-
ing in water that had already been accelerated by the fore flippers. They
might accelerate this water further, but it would be more efficient for
t h e m to work on different water. This is another application of the
principle we have already m e t several times: it is more efficient to get
thrust by accelerating a lot of water to a low velocity, than less water
to a higher velocity.
The long-necked plesiosaurs had extraordinarily long necks, some of
t h e m longer than the whole of the rest of the body. If they s w a m under
water with the long neck stretched out in front it would have been
quite tricky for t h e m to steer a straight course: if the animal acciden-
tally veered slightly to one side, the water, striking the neck obliquely,
would tend to m a k e it veer more. This is the opposite to the effect of
flights on an arrow, w h i c h tend to correct any deviation, pulling the
arrow back to a straight course. T h e difference is that the neck has a
big surface area in front of the animal's center of gravity and the flights
have a big area behind the arrow's one. It seems possible that plesio-
saurs often avoided this problem by s w i m m i n g at the surface with their
necks out of the water. More energy is needed to s w i m at the surface
than to swim well submerged, because an animal at the surface pushes
a bow wave in front of it like a boat, but this might not be a big dis-
advantage if the plesiosaur swam slowly, and in any case it would have
to visit the surface frequently to breathe.
If plesiosaurs had eaten w o r m s or clams, we might suppose that they
138 MARINE REPTILES
used their necks to reach down to the bottom, dabbling like ducks or
swans, but their spiky teeth seem more suitable for catching fishes and
squid-like animals which would probably have been too active to be
caught easily that way. It seems likely that they darted at prey, ex-
tending their long necks to catch things as they swam by. T h e move-
m e n t could have been fast, if the neck was held out of water. Herons
use their long necks to dart at fish, though they stand in the water
instead of floating as plesiosaurs presumably did.
T h e fossil record seems to show that the plesiosaurs became extinct

at the same t i m e as the dinosaurs, 65 million years ago, but some peo-
ple believe that rather similar animals are still living in Loch Ness,
Scotland (figure 9.10). T h e picture is not very like a plesiosaur (notice
the h u m p e d shoulders, and the central ribs in the flippers) but there is
some resemblance.
There have been reports of the Loch Ness m o n s t e r since the Middle
Ages, and t r e m e n d o u s efforts have been made in modern times to get
good evidence of its existence. T h e surface of the lake has been kept
under observation, sonar (echo sounding) has been used, and thousands
of underwater flash photographs have been taken at random in the hope
that the m o n s t e r will swim into view. Some surface photographs have
F I G U R E 9.10. An i m p r e s s i o n of t h e Loch N e s s m o n s t e r , based on e y e w i t n e s s

a c c o u n t s and (unclear) p h o t o g r a p h s . From Scott and Rines 1975. Reprinted by
p e r m i s s i o n . C o p y r i g h t © 1975 M a c m i l l a n M a g a z i n e s Ltd.
MARINE REPTILES 139
been taken that show monster-shaped images, but it never seems cer-
tain that the image is not a floating log or an odd pattern of ripples.
Some sonar traces have detected unexplained objects about 15 meters
long and there are a few hazy underwater photographs (hazy even after
computer enhancement) that show shapes like the neck and flippers
in figure 9.10. If the monster is 15 meters long and has the shape shown
in the picture, it m u s t weigh well over 10 tonnes. If there is one mon-
ster there m u s t be several, or at least there m u s t have been several
until quite recently: no animal is immortal, and very small populations
are in danger of dying out. If there really are monsters there, it seems
odd that we have not got better evidence of their existence.
This chapter has been about three groups of fossil reptiles. The ichth-
yosaurs were beautifully streamlined, like t u n n i e s and dolphins, and
probably swam fast. They may have been e n d o t h e r m s (like tunnies),
they may have dived deep (like dolphins), and they m a y have porpoised
when s w i m m i n g at the surface.
The mosasaurs were giant lizards with flippers instead of legs. Some
have damaged vertebrae that look like s y m p t o m s of the bends, a hazard
of diving.
The plesiosaurs used flippers to swim, probably by underwater flying
rather than rowing. T h e y probably s w a m rather differently from turtles
and penguins, getting thrust only from the downstroke. If so, they must
have been rather slow. Some had remarkably long necks, which may
have been held out of the water and used for darting at prey.
As for the Loch Ness monster, I am not convinced that it exists.
Are you?
Principal Sources
M u c h o f m y i n f o r m a t i o n o n fish s w i m m i n g a n d o n e n d o t h e r m i c fishes c o m e s from

Hoar and Randall (1978). T h e h y p o t h e s i s a b o u t porpoising c o m e s from Au and W e i h s
(1980) and t h e t o p speed for d o l p h i n s from Lang a n d Prior (1966). C l a r k a n d B e m i s
(1979) and D a v e n p o r t a n d o t h e r s (1984) d e s c r i b e h o w p e n g u i n s a n d t u r t l e s s w i m .
R o b i n s o n (1975) a n d Godfrey (1984) d i s c u s s p l e s i o s a u r s w i m m i n g . R o t h s c h i l d a n d
M a r t i n (1987) found t h e e v i d e n c e o f m o s a s a u r s g e t t i n g t h e b e n d s . S c o t t a n d R i n e s
(1975) and W i t c h e l l (1975) p r e s e n t t h e e v i d e n c e for t h e L o c h N e s s m o n s t e r . T h e
o t h e r i t e m s in t h e list are s o u r c e s for i l l u s t r a t i o n s .
Au, D. a n d D. W e i h s . 1980. At h i g h s p e e d s d o l p h i n s s a v e e n e r g y by l e a p i n g . Nature

294:548-550.
Brown, D. S. 1 9 8 1 . T h e E n g l i s h U p p e r Jurassic P l e s i o s a u r o i d e a (Reptilia) a n d a re-
v i e w of t h e p h y l o g e n y a n d classification of t h e P l e s i o s a u r i a . Bulletin of the British
Museum [Natural History) Geology 3 5 : 2 5 3 - 3 4 7 .
140 MARINE REPTILES
C l a r k , B. D. a n d W. B c m i s . 1979. K i n e m a t i c s of s w i m m i n g of p e n g u i n s at t h e De-
t r o i t Z o o . journal of Zoology 1 8 8 : 4 1 1 - 4 2 8 .
D a v e n p o r t , )., S. A. M u n k s , a n d P. J. Oxford. 1984. A c o m p a r i s o n of t h e s w i m m i n g
of m a r i n e a n d f r e s h w a t e r t u r t l e s . Proceedings of the Royal Society B 2 2 0 : 4 4 7 -
475.
Godfrey, S. |. 1984. P l e s i o s a u r s u b a q u e o u s l o c o m o t i o n : a re-appraisal. Neues jahr-
buch fur Ceologie und Paldontologie. Monatshefte. 1984:661-672.
Hoar, W. S. a n d D. J. R a n d a l l . 1978. Fish Physiology. VII: Locomotion. N e w York:
A c a d e m i c Press.
Lang, T. G. and K. Prior. 1966. H y d r o d y n a m i c p e r f o r m a n c e of porpoises [Stenella
attenuata). Science 152:531—533.
R o b i n s o n , |. A. 1975. T h e l o c o m o t i o n of p l e s i o s a u r s . Neues jahrbuch fur Geologie
und Paldontologie, Abhandlungen 149:286-332.
R o m e r , A. S. 1966. Vertebrate Palaeontology. 3d ed. C h i c a g o : U n i v e r s i t y of C h i c a g o
Press.
R o m e r , A. S. 1968. The Procession of Life. L o n d o n : W e i d e n f e l d and N i c h o l s o n .
R o t h s c h i l d , B. a n d L. D. M a r t i n . 1987. A v a s c u l a r n e c r o s i s : o c c u r r e n c e in d i v i n g Cre-
t a c e o u s m o s a s a u r s . Science 2 3 6 : 7 5 - 7 7 .
Scott, P. and R. R i n e s . 1975. N a m i n g t h e Loch N e s s m o n s t e r . Nature 2 5 8 : 4 6 6 - 4 6 7 .
W h e e l e r , A. 1969. The Fishes of the British Isles and North-West Europe. L o n d o n :
Macmillan.
W i t c h e l l , N. 1975. The Loch Ness Story. H a r m o n d s w o r t h : P e n g u i n Books.
X
Death of the Giant Reptiles
A LL T H E magnificent animals that I have been writing about became

extinct at the end of the Cretaceous period, 65 million years ago.
The dinosaurs died out, leaving the birds (which seem to have evolved
from them) as their only descendants. T h e pterosaurs, ichthyosaurs,
mosasaurs, and plesiosaurs died out leaving no descendants. Many in-
vertebrate groups became extinct including the a m m o n i t e s , marine
mollusks with coiled shells which are extremely c o m m o n fossils in
Mesozoic rocks. T h e extinctions were devastating, but there were also
a lot of survivals. T h e m a m m a l s (which had all been small in the time
of the dinosaurs) survived well, and so did the land plants. Most of the
main groups of lizards (other than mosasaurs), snakes, turtles and croc-
odiles also survived. What killed the dinosaurs and left the crocodiles?
There have been a lot of suggestions about w h y the dinosaurs w e n t
extinct, some of t h e m distinctly far-fetched. T w o hypotheses are strongly
supported at present, by different groups of scientists. O n e says that
the earth was hit by a collossal meteorite (a l u m p of material from
outer space) and the other says that there was a period of violent vol-
canic activity.
The meteorite hypothesis started w i t h a strange observation. T h e
latest Cretaceous rocks in central Italy and the earliest rocks of the
next period (the Tertiary) are both limestones. Each contains fossil shells
of foraminiferans (microscopic marine animals) typical of its period.
Between these limestone layers is a layer of clay, two centimeters thick,
with no fossils in it. A team of scientists led by Drs. Luis and Walter
Alvarez (father and son) analyzed this clay using n e u t r o n activation
analysis, a technique that can measure tiny quantities of rare elements.
They found remarkably high concentrations of iridium, one of the plat-
i n u m group of metals.
When I say high, I mean 9 parts per billion (9 parts in 10 : I am using
y
142 DEATH OF THE G I A N T REPTILES
the American billion, not the larger British one). T h a t m a y seem too
little to get excited about, but it is 30 times higher than in the lime-
stone immediately below or a short distance above (figure 10.1). Iridi-
um concentrations are generally exceedingly low in the earth's crust
but m u c h higher in meteorites, typically 500 parts per billion. Could
the iridium in the clay have come from a meteorite?
After the Italian rocks had been analyzed, samples of rock were taken
from other places, scattered around the world, where Cretaceous and
Tertiary deposits meet w i t h no apparent interruption. These were ana-
lyzed in the same way, and high iridium concentrations were found in
t h e m all. T h e iridium layer seemed to be everywhere.
A meteorite hitting the earth might explode, scattering iridium-rich
F I G U R E 10.1. A graph s h o w i n g i r i d i u m c o n c e n t r a t i o n s in the clay at t h e Cre-

t a c e o u s - T e r t i a r y b o u n d a r y at G u b b i o , Italy, and in t h e l i m e s t o n e s i m m e d i a t e l y
a b o v e a n d b e l o w it. T h e c a l c i u m c a r b o n a t e w a s d i s s o l v e d o u t w i t h a c i d b e f o r e
t h e a n a l y s e s w e r e c a r r i e d o u t , a n d t h e c o n c e n t r a t i o n s refer t o t h e a c i d - i n s o l u b l e
r e s i d u e . T h e d a t a a r e f r o m L . W . A l v a r e z e t al., Science (1980), 2 0 8 : 1 0 9 5 - 1 1 0 8 .
DEATH OF THE GIANT REPTILES 143
dust, but any ordinary explosion would scatter material only over a
restricted area. Here we have material scattered all over the world. T h e
Alvarez team suggested that a huge meteorite might have disintegrated
in a collossal explosion, throwing dust m a n y kilometers up into the
atmosphere. If this dust were fine enough it would be slow to settle,
and might get scattered all over the earth.
It is quite easy to calculate how big the meteorite would have had
to be, to scatter so m u c h iridium. First we need to know h o w m u c h
extra iridium there is, above w h a t would be expected in the same
thickness of ordinary rock. Analyses from twenty-one widely scattered
places give an average of 0.6 milligrams of extra iridium per square
meter of the earth's surface. T h e area of the earth is 5 x 1 0 (500 14
million million] square meters, so the total a m o u n t of iridium can be

estimated as 0.6 x 5 x 1 0 = 3 x 1 0 milligrams or 300,000 tonnes.
14 14
T h e meteorite would probably have contained about 500 parts per bil-
lion of iridium (1 part in 2 million), so we m u s t multiply the 300,000
tonnes by 2 million to get an estimate for the total m a s s of the me-
teorite, 600 billion tonnes.
Typical meteorites have densities of about 2.2 tonnes per cubic me-
ter (which is rather lower than m o s t other rocks) so a 600-billion-tonne
one would have a volume of 270 billion cubic meters. A sphere of that
volume would have a diameter of 9 kilometers. It would be similar in
size to M a n h a t t a n Island (which is about 20 kilometers long and 4 kilo-
meters wide).
It may seem far-fetched to imagine such a thing hitting the earth,
but it is not too improbable. As well as planets orbiting the sun, there
are a lot of smaller bodies called asteroids, a few kilometers in diam-
eter. The meteorites that have been observed landing on earth seem to
be fragments from collisions between asteroids, but there is a constant
danger of whole asteroids hitting us. It has been estimated from tele-
scope observations that there are about a thousand, w i t h diameters of
a kilometer or more, whose orbits take t h e m inside the earth's orbit
at times and outside it at others. N o n e of these asteroids have collided
with the earth in historic t i m e s (so far we have been lucky), but even-
tually some will. A few craters have been found that are believed to
have been made in the distant past by small asteroids, and it has been
calculated that the earth is likely to be hit by an asteroid of 10 kilo-
meters or more diameter about once every 100 million years. T h a t is
very seldom, but the event we are trying to explain happened just once,
65 million years ago.
T h e earth, traveling its orbit round the sun, is hurtling through space
at 30 kilometers per second (one hundred times the speed of sound in
air). Asteroids travel round the sun in the same direction, so there will
be no head-on collisions. Figure 10.2 shows how the earth, traveling

its near-circular orbit, might be hit by an asteroid with a more ellip-
tical orbit. Astronomers tell us that the speed of an approaching as-
teroid, relative to the earth, would probably be something like 20 kilo-
meters per second.
We will calculate the energy of an impact at this speed. This is the
kinetic energy of the asteroid, due to its movement relative to the earth.
Kinetic energy is j (mass) x (speed) , but in using the formula we m u s t
2
be careful about units. Six hundred billion tonnes is 600 million mil-
lion kilograms. T w e n t y kilometers per second is 20,000 meters per sec-
ond. If we put into the formula the mass in kilograms and the speed
in meters per second we get the energy in joules; it is about 10 ' (100,000 1
million million million) joules. T h i s is e n o r m o u s — e q u i v a l e n t to the

explosion of 60 million megatonnes of T N T . The atomic bombs dropped
in Japan had energies of only 0.02 megatonnes each. The biggest ex-
plosion of modern times, the explosion of the island volcano Krakatoa
in 1883, had about 200 megatonnes energy.
My calculation is very rough because the speed that I used in it could
be badly wrong, but it seems clear that the landing of the asteroid would
have been incomparably more devastating than anything people have
ever experienced. T h e energy would have been amply sufficient to va-
porize the asteroid. T w e n t y million joules are needed to vaporize a ki-
logram of rock so 600 million million kilograms could be vaporized by
FIGURE 10.2. Collision b e t w e e n the earth and an asteroid. N o t to scale.

12,000 million million million joules (1.2 x 1 0 joules). This is only

22
one-eighth of the estimated kinetic energy.

If the asteroid had hit dry land it would have made an e n o r m o u s
hole, but no crater that could have been made by it has been found.
This need not worry us: it would be more likely to land in the oceans
that cover 70 percent of the earth's surface. If it did fall in an ocean it
would m a k e less of a crater, and even a big crater in the ocean floor
would be hard to find.
A lot of the energy of the asteroid would be transmitted to the ocean
water, turning some of it to high-pressure steam, but there would be
enough left to vaporize most of the meteorite. There would be a terrific
explosion that would blast a column of steam and rock vapour high
into the atmosphere where they would condense out as tiny ice crys-
tals and dust particles. T h e dust would sink down again onto the earth
but the rate of settling would depend on the size of the particles: it
would probably take a few m o n t h s .
N o w we will think about how the catastrophe could have affected
dinosaurs and other animals. First, the dust in the atmosphere would
have blotted out the sun. Sunlight all over the world is d i m m e d by
dust after major volcanic eruptions, and would be d i m m e d far more by
the catastrophe we are imagining. It has been estimated that after the
Krakatoa eruption, sunlight was d i m m e d by 3 percent to 0.97 of its
normal intensity. Twice as m u c h dust would dim it to (0.97) times 2
the usual intensity, three times as m u c h to (0.97)' and so on. If the

asteroid threw up 200 t i m e s as m u c h dust as Krakatoa (and the ratio
of energies suggests it would have t h r o w n up far more than that) sun-
light would be d i m m e d to (0.97) "" - 0.002 t i m e s its normal intensity:
2
the world would have been plunged in darkness.

Calculations of the size of the dust particles and of the rate at which
they would settle suggest that the darkness would have lasted for sev-
eral m o n t h s . Plants would have suffered because they depend on the
energy of sunlight to m a k e foodstuffs by the process of photosynthesis.
Many land plants would probably have survived a few m o n t h s darkness
(remember that deciduous trees lose their leaves each fall and have to
survive a few m o n t h s w i t h o u t photosynthesis), and other land plants
would probably have survived as seeds. However, the microscopic plants
that float as plankton in lakes and seas would probably have suffered
badly, because they are too small to have substantial food reserves.
Almost all the animals in lakes and seas get their energy ultimately
from these microscopic plants: tiny crustaceans and other animal
plankton eat the plants and are in turn eaten by fish and other larger
animals. Life in lakes and seas would suffer very badly.
A second effect would result from the blotting out of sunlight: the
146 DEATH OF THE GIANT REPTILES
earth's surface would get cold. T h e effect on the oceans would not be
great because of their e n o r m o u s heat capacity, but there would be se-
vere frosts on land that might be lethal to many plants and animals.
A third possible effect would be acid rain. T h e high temperatures of
the explosion would m a k e some of the nitrogen in the atmosphere
combine with oxygen to form nitrogen oxides. These would react with
water and more oxygen to form nitric acid, which would fall from the
atmosphere in rain.
Acid rain due to very different causes is a serious modern problem.
Nitrogen oxides are formed in the engines of m o t o r vehicles and re-
leased into the atmosphere w i t h the exhaust. Sulphur dioxide is emit-
ted by coal-burning power stations. T h e nitrogen and sulphur oxides
react w i t h oxygen and water to form nitric and sulphuric acids, which
fall in rain. T h e acid rain falling on trees m a k e s leaves yellow and fall
off. When it drains into lakes it m a k e s t h e m acid, sometimes too acid
for fish to survive. Many forests in industrial countries are in poor health
and m a n y lakes have lost their stocks of fish. T h e acid rain after the
asteroid explosion would have had similar effects.
T h e asteroid explosion would have had disastrous effects on many
kinds of animals and plants. Nevertheless, the asteroid hypothesis for
the extinctions at the end of the Cretaceous has several problems. One
is that the high iridium concentrations are not limited to very thin
layers of rock as would be expected if they had been formed by dust
settling in the few m o n t h s after the explosion. Instead, they extend
through thicknesses of 30 to 100 centimeters, that probably took sev-
eral tens of thousands of years to form. Indeed, some American sam-
ples show several iridium-rich layers sandwiched between iridium-poor
ones. Another problem is that the extinctions do not seem to have
happened all at once. T h e n u m b e r s of a m m o n i t e and dinosaur species
seem to have declined gradually during the last few million years of
the Cretaceous, and the last N o r t h American dinosaurs are in rocks
above the iridium-rich layer, formed 40,000 years after it.
These observations seem to favor the volcanic hypothesis, which
postulates a few tens of thousands of years of intense volcanic activity.
Volcanoes throw up material from deep inside the earth, where iridium
concentrations are m u c h higher than in surface rocks, though lower
than in meteorites. The iridium-rich layers may have come from vol-
canoes.
Major eruptions throw enough dust into the upper atmosphere to
dim sunlight perceptibly, but eruptions big enough to black out the
sun and stop photosynthesis seem unlikely. Volcanoes seem more likely
to cause extinctions by way of acid rain. They emit sulphur dioxide
and other gases as well as m o l t e n rock. T h e sulphur dioxide (like the
DEATH OF THE G I A N T REPTILES 147
sulphur dioxide from power stations) eventually falls in rain as sul-

phuric acid. Scientists have analyzed emissions from a Hawaiian vol-
cano, measuring the quantities of sulphur dioxide and iridium. Their
m e a s u r e m e n t s suggest that if the 300,000 tonnes of iridium in the irid-
ium-rich layers came from volcanoes, they would have been accom-
panied by about 10 million million tonnes of sulphur dioxide. If this
was emitted over a period of 10,000 to 100,000 years, the average rate
of sulphur dioxide emission would have been between 100 million and
1 billion tonnes per year. The rate would probably vary, with peak rates
far above average.
T h e surphur dioxide, released annually in the United States and Eu-
rope by burning coal and oil, totals 80 million tonnes, and the nitrogen
oxide emissions are less. I have already described the damage that these
emissions are causing, through acid rain. T h e acid rain in the worst
parts of the supposed period of volcanic activity would have been far
worse.
It would have been so m u c h worse that the oceans would have been
seriously affected, as well as lakes. At present the oceans are slightly
alkaline with a pH value of 8.2, but the acid from 10 million million
tonnes of sulphur dioxide would reduce the pH to about 7.4. T h i s is
still very slightly on the alkaline side of neutrality (pH7 is neutral and
anything less is acid), but it is not alkaline enough for foraminiferans.
It is also near the limit for another group of microscopic plankton, the
coccoliths.
Foraminiferans have calcium carbonate shells and coccoliths have
plates of calcium carbonate on their outer surfaces, and calcium car-
bonate dissolves in acid. Foraminiferans need a pH of 7.6 or more and
coccoliths need at least 7.0-7.3. Both groups suffered m a n y extinctions
at the end of the Cretaceous but dinoflagellates and other groups of
plankton, w i t h o u t calcium carbonate skeletons, survived better.
Ozone is a form of oxygen that is rare at ground level but relatively
more plentiful in a layer high in the atmosphere. It absorbs m u c h of
the ultraviolet radiation from the sun, protecting living things from
these harmful rays. T h e ozone layer is depleted after major eruptions
because volcanoes inject a little hydrochloric acid into the atmosphere,
as well as the m u c h larger quantity of sulphuric acid. T h e hydrochloric
acid reacts with the ozone in the ozone layer to form chlorine, water,
and ordinary oxygen. It has been calculated that 8 percent of the ozone
was destroyed after the Krakatoa eruption. T h e m a n y eruptions that
are supposed to have happened at the end of the Cretaceous would
have nearly destroyed the ozone layer, leaving animals and plants ex-
posed to abnormally high doses of ultraviolet radiation. This might have
been fatal for m a n y of them, but the m a m m a l s of the time were small
and would have survived if they spent their days in burrows and were
active mainly at night.
T h e volcanic hypothesis seems quite attractive but there is at least
one observation that it seems unable to explain. Damaged sand grains
have been found wherever they have been looked for, in samples from
the iridium-rich layer from various places, scattered around the world.
Sand grains are small quartz crystals, made of neatly stacked layers of
atoms. T h e damaged ones look cracked (when examined under the mi-
croscope) because some of their layers of a t o m s have been thrown into
disarray. Similar damage is found in sand grains from meteorite craters
and is believed to have been caused by shock waves. T h e volcanic hy-
pothesis seems unable to explain shocked quartz being scattered widely
round the world.
Both hypotheses seem reasonably plausible. Collision of an asteroid
with the earth, and a prolonged period of fierce volcanic activity, would
each have had dire consequences, and would probably have caused
widespread extinctions. T h e volcanic hypothesis is possibly the better
of the two, in explaining why the extinctions were so selective: acid
rain would kill foraminiferans but not dinoflagellates, and ultraviolet
radiation would be more damaging for diurnal dinosaurs than for noc-
turnal m a m m a l s . However, the asteroid hypothesis seems better able
to explain the shocked quartz. T h e supporters of the two hypotheses
are still arguing fiercely, and it is quite possible that neither is right.
Indeed, it has recently been suggested that the extinctions were due to
another cause, a shower of comets hitting the earth. The effects of comet
impacts would be m u c h like those of asteroid impacts: comets are
massive bodies travelling at exceedingly high speeds, and contain more
iridium than the surface rocks of the earth. A series of comet impacts
could explain extinctions spaced out in time, and there are theoretical
reasons for expecting comets to come in showers lasting about a mil-
lion years. T h e implications of the comet shower hypothesis have not
yet been worked out in as m u c h detail as those of the asteroid and
volcano hypotheses. It remains to be seen which (if any) of the three
hypotheses t r i u m p h s .
Principal Sources
T h e m e t e o r i t e h y p o t h e s e s w a s p u t forward b y A l v a r e z c t al. (19801 and d i s c u s s e d i n

m u c h m o r e d e t a i l b y Silver a n d S c h u l t z (1982). T h e v o l c a n o h y p o t h e s i s h a s b e e n
p r e s e n t e d by Officer et al. (1987). Sloan et al. (1986) p u b l i s h e d e v i d e n c e t h a t t h e
d i n o s a u r s declined gradually. Bohor, M o d r e s k i , and Foord (1987) presented t h e shocked
q u a r t z e v i d e n c e . H u t a n d o t h e r s (1987) p r e s e n t e d t h e c o m e t s h o w e r h y p o t h e s i s .
Alvarez, L. W . W. Alvarez, F. Asaro, a n d H. V. M i c h e l . 1980. E x t r a t e r r e s t r i a l c a u s e

:
for t h e C r e t a c e o u s - T e r t i a r y e x t i n c t i o n . Science 2 0 8 : 1 0 9 5 - 1 1 0 8 .
Bohor, B. F., P. ). M o d r e s k i , and E. E. Foord. 1987. S h o c k e d q u a r t z in t h e C r e t a c e o u s -
T e r t i a r y b o u n d a r y c l a y s : E v i d e n c e for a global d i s t r i b u t i o n . Science 2 3 6 : 7 0 5 - 7 0 9 .
H u t , P. and o t h e r s (1987). C o m e t s h o w e r s as a c a u s e of m a s s e x t i n c t i o n s . Nature
329:118-126.
Officer, C. B., A. H a l l a m , C. L. D r a k e , a n d J. D. D e v i n c . 1987. L a t e C r e t a c e o u s and
p a r o x y s m a l C r e t a c e o u s / T e r t i a r y e x t i n c t i o n s . Nature 3 2 6 : 1 4 3 - 1 4 9 .
Silver, L. T. a n d P. H. S c h u l t z . eds. 1982. G e o l o g i c a l i m p l i c a t i o n s of i m p a c t s of large
a s t e r o i d s a n d c o m e t s on t h e e a r t h . Geological Society of America Special Paper
190:1-528.
Sloan, R. E., J. K. Rigby, L. M. Van Valen, a n d D. G a b r i e l . 1986. G r a d u a l d i n o s a u r
e x t i n c t i o n and s i m u l t a n e o u s u n g u l a t e r a d i a t i o n i n t h e H e l l C r e e k F o r m a t i o n . Sci-
ence 2 3 2 : 6 2 9 - 6 3 3 .
XI
Giant Birds
A RCHAEOPTERYX, THE oldest k n o w n bird, lived in the Jurassic pe-

riod, quite early in the t i m e of the dinosaurs. It was only about
the size of a magpie and no giant birds appeared until the Tertiary, after
the dinosaurs had gone.
T h e teratorns were birds of prey that lived in America then. Figure
11.1 will give you an impression of their size: (b) shows the wing bones
of a Californian condor, one of the largest modern birds of prey; (d)
shows the wing bones of the Wandering albatross, which has the big-
gest wing span of all modern birds; (c) shows the wing skeleton of the
best-known teratorn, Teratornis merriami: it is almost as long as the
albatross wing skeleton, and rather stouter; finally, (a) is amazing. It
is the h u m e r o u s (the bone from the base of the wing) of Argentavis
magnificens, the biggest k n o w n teratorn. It is more than twice as long
as the condor h u m e r u s shown immediately below it.
Only one skeleton of Argentavis has been found (in Argentina). The
wing bones are incomplete but the pieces that have been found are
enough to show that the wings were strong and well developed. It seems
impossible to avoid the conclusion that those huge wings were for use,
that Argentavis could fly although it was far bigger than any modern
flying bird.
T h e wing span of the condor is ten times the length of the h u m e r u s .
If Argentavis was built to the same proportions its span was about 6
meters, far more than the spans of the condor (2.7 meters) or albatross
(3.4 meters). Its biggest wing feathers m u s t have been about 1.5 meters
long and 200 millimeters wide. Its body mass has been estimated from
the circumference of a leg bone in the same way as was done for di-
nosaurs, using a graph like Figure 2.5 showing tibiotarsus circumfer-
ence and body mass for 324 species of modern birds. The body masses
corresponding to the tibiotarsus circumferences of teratorns were read
G I A N T BIRDS 151
F I G U R E 1 1 . 1 . W i n g b o n e s of (a] Argentavis, t h e l a r g e s t t e r a t o m ; (b) a C a l i f o r -

n i a n c o n d o r ; (c) a W a n d e r i n g a l b a t r o s s ; a n d (d) Teratornis. F r o m C a m p b e l l a n d
T o n n i 1983.
off from the graph: 14 kilograms for Teratornis merriami and 80 ki-
lograms (heavier than most men) for Argentavis.
That 80 kilograms is the best estimate that has been made of the
mass of Argentavis, but there is a lot of uncertainty about it. Birds of
equal mass, of different species, m a y have considerably different bone
thicknesses. Statistical analysis of the data tells us that the conclusion
should be no more precise than this: there is 95 percent probability
(the odds are 19 to 1) that the mass of Argentavis lay between 37 and
166 kilograms. Even w i t h that m u c h uncertainty it is clear that Ar-
gentavis was much heavier than the Californian condor (about 10 ki-
lograms) or even the Kori bustard, which reaches about 16 kilograms
and seems to be the heaviest modern flying bird.
Argentavis was also m u c h heavier than Pteranodon, the giant ptero-
saur (figure 8.2), which was so lightly built that its m a s s seems to have
been no more than about 15 kilograms. However, Pteranodon had the
larger wing span (7 meters. We estimated the span of Argentavis as
only 6 meters). T h e few fragments that have been found of the even
larger pterosaur, Quetzalcoatlus, suggest a span of about 12 meters and
a mass of possibly about 60 kilograms. Argentavis and Quetzalcoatlus
are rivals for the title of the biggest flying animal of all time.
152 GIANT BIRDS
T h e largest modern birds cannot fly . They are the ostrich (up to 120
kg), cassowaries (60 kg), the e m u (50 kg) and the Emperor penguin (40
kg). With the exception of the penguin, these are members of the group
called the ratites, which also includes the rheas and kiwis.
Ostriches and other ratites are like enormously overgrown chicks.
They have tiny wings, useless for flight, and well-developed legs. They
have fluffy plumage instead of the blade-like feathers of other adult
birds. T h e y also have some chick-like features in their skeletons. They
are believed to have evolved by processes of development getting out
of step w i t h each other: they grow large and sexually mature while
keeping a lot of juvenile features.
T h e biggest extinct birds are also ratites. They are the moas, which
lived in N e w Zealand, and the elephant birds, in Madagascar. The big-
gest moa is Dinornis maximus, 3.5 meters tall (twice the height of an
average man, figure 11.2). T h e biggest elephant birds looked very sim-
ilar and were about 3 meters tall. There were also some giant birds
that were not ratites. Diatryma, a wicked-looking predator that lived
in N o r t h America, was about 2 meters tall (figure 11.2). It lived quite
early in the Cenozoic era but the moas and elephant birds are more
recent. Indeed, the moas survived in New Zealand until after the Maoris
arrived.
I have made scale models of moas and used t h e m to estimate the
FIGURE 11.2. Dinornis maximus (the largest moa), Diatryma steini, and an
a d u l t m a n , all t o t h e s a m e scale.
GIANT BIRDS 1 S3
masses of the living birds, in m u c h the same way as I estimated the

masses of dinosaurs (chapger 2). I modeled the m a i n features of the
skeleton in wire and then added clay to represent the flesh, m a k i n g
models that represented the birds as if they had been plucked. I mea-
sured the volumes of the models and used t h e m to calculate the masses
of the plucked birds, assuming that their densities were the same as
the density of a plucked goose carcase (which I measured). Finally, I
added an allowance for the feathers, w h i c h I assumed to be the same
fraction of body mass as in turkeys and kiwis. T h e result for a big
Dinornis was 240 kilograms, about the same as a large tiger. T h e big-
gest elephant birds were stouter, although they were a little shorter,
and may have been nearly twice as heavy.
Though Dinornis was the biggest moa, the one that fascinates me is
Pachyornis elephantopus, shown in figure 11.3. Its splendid name means
"fat bird with elephant's feet," and seems very suitable. Its leg bones
are amazingly thick. Compare it w i t h the ostrich, drawn beside it to
the same scale. I estimate this moa's mass as 130 kilograms (from mea-
surements on a model) and the ostrich, whose skeleton is illustrated,
as only 68 kilograms, but even so the moa bones look disproportion-
ately thick.
Appearances can be deceptive, so I measured the m o a ' s bones and
calculated strength indicators in the same way as for dinosaurs (chapter
4). The results are shown in table 11.1. T h e value for the tibiotarsus
(shin bone) is about the same as for an ostrich, and those for the other
two bones are twice as high as for the ostrich. To be consistent w i t h
my line of argument in chapter 4, I should conclude that Pachyornis
was at least as athletic as ostriches, but I find that hard to believe.
Ostriches are exceedingly fast runners, probably faster than the African
antelopes. Should I conclude that moas were also exceedingly fast? I
cannot believe that they were, w i t h those clumsy-looking legs.
The key to the problem may be that moas seem to have had no need
to run. They fed on plants, as remains of their stomach c o n t e n t s show,
and so had no need to run to capture food. There seem to have been
no big predators in N e w Zealand, until the Maoris arrived, so there
was nothing to run away from. (That is to say, there was nothing to
run away from while they were evolving. T h e y seem to have been easy
prey to the Maoris, w h o h u n t e d t h e m to extinction.) It was not the
same for the dinosaurs: the flesh-eating dinosaurs had to run to catch
prey and the plant-eating ones had to run to escape. This difference
between moas and dinosaurs may justify a different interpretation of
their strength indicators.
My idea involves safety factors. Suppose an engineer is designing a
small bridge to carry a m a x i m u m load of ten tonnes. He would be an
154 GIANT BIRDS
optimist if he calculated the thickness of steel that could just support

ten tonnes w i t h o u t breaking, and ordered steel that thick. Any repu-
table engineer would allow a safety factor: he might design the bridge
to be able to carry t w e n t y tonnes although he expected the m a x i m u m
load to be only ten. T h e reason for this is that neither load nor strength
can be predicted precisely. An unexpectedly large load may arrive, or
the steel may be substandard, and in either case a bridge that was ex-
pected to be strong enough m a y fail. T h e bigger the safety factor the
less likely this is to happen.
F I G U R E 1 1 . 3 . S k e l e t o n s o f (a| Pachyornis a n d (b) a n o s t r i c h , t o t h e s a m e s c a l e .

F r o m A l e x a n d e r 1983a.
GIANT BIRDS 155
TABLE 11.1. Strength indicators [Z/W\, see p. 53) for leg bones of
an ostrich and the moa Pachyornis elephantopus.
Strength indicator (square meters per giganewton) for:
Femur Tibiotarsus Tarsometatarsus
ostrich 45 18 17
moa 94 17 39
A stronger bridge is less likely to fail but costs more to build. Beyond
a certain point, the slight advantage of extra safety obtained by m a k i n g
it stronger still is not worth the extra cost. T h e ideal strength depends
on the cost of the materials. If we had to m a k e bridges of p l a t i n u m we
would make t h e m weaker and live more dangerously. If cheap second-
hand steel were available we might m a k e a bridge extra strong.
The evolution of skeletons has also involved balancing safety against
cost. In this case the cost is partly the cost in energy and materials of
growing a stronger bone, but is largely the penalty of having to carry
extra bone around. Thick leg bones may be less likely to break in a
fall, but they make it harder to run fast, just as people find it hard to
sprint in heavy boots. M e a s u r e m e n t s and calculations on ostriches and
several kinds of m a m m a l have shown that their leg bones are about
three times as strong as is necessary to withstand the forces involved
in strenuous activities such as running and jumping. These bones are
built to safety factors of about three, and so can stand m a n y of the
larger forces that occur accidentally, for example in falls and collisions.
Moas may have had larger safety factors. However strong their bones,
there would always be some danger of an accident bad enough to break
them. If they had no need to run, they might not be inconvenienced
much by heavy bones. T h e cost of extra strength might be less for t h e m
than for ostriches, which have to run to escape from lions. If strength
were cheap, ideal safety factors would be high, w h i c h may explain the
remarkably thick leg bones of Pachyornis and (to a lesser extent) other
moas.
Several sets of moa footprints have been found and I have calculated
speeds for them, in the same way as for dinosaurs (chapter 3). All of
t h e m seem to have been moving between 0.8 and 2.0 meters per sec-
ond, which would have been walking speeds. This does not prove that
they never ran, but at least it does not contradict the suggestion that
they were not very athletic.
156 GIANT BIRDS
My belief, that moa leg bones had high safety factors, has been chal-
lenged. Palaeontologists in Tubingen have suggested that moas may
have lived in thick undergrowth and may have needed very strong legs
to force their way through. I find that hard to believe. An animal that
behaved like a bulldozer would use a lot more energy than one that
slipped through small gaps, or avoided the densest patches of vegeta-
tion, and might be a poor competitor. Nevertheless, the possibility should
be considered.
T h e legs of elephant birds are little less remarkable than the legs of
moas, but it is their eggs that I w a n t to write about, the biggest of all
k n o w n eggs. Quite a lot of elephant-bird egg shells have been found in
Madagascar, some with the bones of embryos still inside them. The
eggs of the biggest species are 30 centimeters long with a volume of 9
liters (2.4 U.S. gallons). Ostrich eggs are only about half as long, with
a volume of 1.3 liters, and the eggs of all modern reptiles are m u c h
smaller. Even k n o w n dinosaur eggs are smaller than elephant bird eggs:
the biggest I know of are only 25 centimeters long.
Let us t h i n k what problems there might be, for very large eggs. First
there is the problem of ventilation. Bird embryos are not hermetically
sealed in their eggs, like cans of soup. T h e eggshell is porous, allowing
gases to diffuse in and out. This enables the embryo to get the oxygen
it needs for respiration, and to get rid of waste carbon dioxide.
T h i n k of two eggs, one twice the length of the other. It has eight
times the volume of the smaller egg, and the embryo in it, just before
hatching (when it uses oxygen fastest), is eight times as heavy. The big
embryo uses oxygen faster than the small one, but not eight times as
fast, because rates of oxygen c o n s u m p t i o n are not strictly proportional
to body mass either for adult animals (figure 7.1) or for embryos. The
large embryo will probably use oxygen only four or five times as fast
as the small one.
T h e more pores there are, or the wider the pores, the faster oxygen
can diffuse in. However, the thicker the shell, the further the oxygen
has to diffuse and the lower the rate of diffusion. T h e m a x i m u m rate
of diffusion that a shell allows is proportional to
number of pores x area of each pore

-------------------------------------------
thickness of shell.
If the two eggs were precise scale models of each other they would have
equal n u m b e r s of pores and the larger one would have pores of twice
the diameter, therefore four t i m e s the cross-sectional area, but its shell
would be twice as thick. It would allow oxygen to diffuse just twice
as fast but the embryo in it would need oxygen, as we have seen, four
GIANT BIRDS 157
or five times as fast. This tells us that big eggs need more porous shells
than small ones. An excessively big egg would need a shell so riddled
with pores as to be seriously weakened. If this shell were made thicker,
to strengthen it, it would have to be more porous still. T h e need to be
sufficiently porous m u s t set an upper limit to the sizes of eggs.
Even elephant bird eggs are probably a long way from that limit.
Chicken eggs have very sparse pores, piercing only 0.02 percent of the
are of the shell. Ostrich eggs have to be m u c h more porous, w i t h 0.2
percent of their area accounted for by pores. Elephant bird eggs m u s t
have been more porous still, but even if the pores were 2 percent of
their area the shells would not be seriously weakened. I know no mea-
surements of their pores so I cannot state the exact percentage.
Elephant bird eggs may be near an upper size limit, for a different
reason. Eggs have to be strong enough to withstand the forces that act
on them, when the parent birds get on and off the nest, but they m u s t
be weak enough for the hatching chick to break its way out. T h i n k
again of two eggs, one twice as long as the other and eight t i m e s as
heavy. It probably needs to be more than eight t i m e s as strong. T h i s
is because big birds are heavier, relative to the masses of their eggs,
than small ones: very small birds are about five t i m e s as heavy as their
eggs but ostriches are fifty t i m e s as heavy as their eggs. T h e big egg
seems to need to be more than eight t i m e s as strong, but if its shell is
just twice as thick, it will only be four t i m e s as strong. (It is a general
rule for objects of the same shape, made of the same material, that
strength is proportional to (length) ). This m e a n s that bigger eggs need
2
relatively thicker shells. An egg that is twice as long as another is gen-

erally found to have a shell about three t i m e s as thick. T h e eggshell
is 4 percent of the mass of a h u m m i n g b i r d egg but 17 percent of the
mass of an ostrich egg.
Hatching chicks break their eggs open by h a m m e r i n g at the shell.
When you break something by h a m m e r i n g or by any other kind of im-
pact, what decides w h e t h e r it breaks or not is the energy of the blow.
A heavy h a m m e r brought down fast has more kinetic energy than a
light h a m m e r moving slowly, and is more likely to break things. The-
ory tells us that the energies needed to break egg shells should be about
proportional to the masses of the shells. T h e energies that chicks can
put into blows should be about proportional to the masses of the chicks
and so to the masses of the egg contents (since the hatching chick fills
the shell). If bigger eggs have relatively thicker shells, it will be harder
for their chicks to break out of them. Too big an egg would be an un-
breakable prison.
One final thought about elephant birds: they m u s t have been blessed
with patience. Small egg hatch soon but big ones take longer. Very
158 GIANT BIRDS
small bird eggs hatch in about 15 days, and ostrich and e m u eggs take
about 50 days. If the trend continues, elephant bird eggs would have
taken about 90 days to hatch.
This chapter has been about three groups of giant birds: the teratorns,
moas, and elephant birds. T h e teratorns had well-developed wings and
could presumably fly, though the biggest seem to have been five times
as heavy as any modern flying bird.
T h e moas and elephant birds were like outsize ostriches and could
not fly. Some moas had astonishingly thick leg bones which seem un-
necessarily strong, for animals that do not look very athletic. I suggest
that they m a y have evolved unusually high safety factors because extra
m a s s in the legs would be little disadvantage, if moas did not have to
run. T h e y were not threatened by any predator until h u m a n s arrived
in N e w Zealand.
Elephant birds laid the biggest k n o w n eggs. These needed thick shells
to protect t h e m from damage by the parents, but thick shells need to
be very porous (to let oxygen diffuse in fast enough) and are difficult
for the hatching chick to break out of. Elephant bird eggs may have
been near the limit of size set by the difficulty of hatching.
Principal Sources
C a m p b e l l a n d T o n n i (1983) d i s c u s s e d t h e flying a b i l i t y o f t e r a t o r n s . T h e d i s c u s s i o n
of m o a s is based on t w o p a p e r s of my o w n , o n e (1983) on Pachyornis a n d o n e (1981)
o n safety factors i n a n i m a l s k e l e t o n s g e n e r a l l y . M o s t o f m y i n f o r m a t i o n a b o u t egg-
shells c o m e s from A n d e r s o n , Rahn, and Prange (1979), R a h n , Ar, and Paganclh (1979),
a n d T u l l e t t (1984).
A l e x a n d e r , R. M c N . 1 9 8 1 . F a c t o r s of safety in t h e s t r u c t u r e of a n i m a l s . Science Prog-

ress 6 7 : 1 0 9 - 1 3 0 .
A l e x a n d e r , R. M c N . 1983a. A l l o m e t r y of t h e leg b o n e s of m o a s ( D i n o r n i t h c s ) and
o t h e r b i r d s , journal of Zoology 2 0 0 : 2 1 5 - 2 3 1 .
A l e x a n d e r , R. M c N . 1983b. On t h e m a s s i v e legs of a m o a [Pachyornis elephanlopus,
D i n o r n i t h e s ) . fournal of Zoology 2 0 1 : 3 6 3 : 3 7 6 .
A n d e r s o n , J. F., H. R a h n , a n d H. D. P r a n g e . 1979. S c a l i n g of s u p p o r t i v e t i s s u e m a s s .
Quarterly Review of Biology 5 4 : 1 3 9 - 1 4 8 .
C a m p b e l l , K. E. a n d E. P. T o n n i . 1983. Size a n d l o c o m o t i o n in t e r a t o r n s (Aves: Ter-
a t o r n i t h i d a e ) . The Auk 1 0 0 : 3 9 0 - 4 0 3 .
R a h n , H., A. Ar, a n d C. V. P a g a n e l l i . 1979. H o w bird eggs b r e a t h e . Scientific Amer-
ican 240(2|:38-47.
T u l l e t t , S. G. 1984. T h e p o r o s i t y of a v i a n e g g s h e l l s . Comparative Biochemistry and
Physiology 78A:5-13.
Reif, W. E. and H. S y l i n - R o b e r t s . 1987. On t h e r o b u s t n e s s of m o a leg b o n e s . Neues
jahrbuch fur Geologie and Paldontologie, Monatshefte 1987:155-160.
XII
Giant Mammals
T HERE WERE no big m a m m a l s while the dinosaurs lived but m a n y

m a m m a l s of elephant or rhinoceros size evolved during the Ceno-
zoic era.
The best k n o w n of the giant extinct m a m m a l s were the mastodons
and m a m m o t h s . Mastodons are primitive elephants, distinguished by
their simple teeth. M a m m o t h s are m u c h more like modern elephants.
Both survived to overlap in t i m e with h u m a n s , and m a n y cave paint-
ings of m a m m o t h s have been found.
The biggest m a m m o t h species [Mammuthus imperator of N o r t h
America) stood 4 meters tall at the shoulder. Very large (6 tonne) Af-
rican elephants are only 3.3 meters tall. T h e m a m m o u t h , 1.2 t i m e s as
tall, must have been about 1.2 t i m e s as heavy, about 10 tonnes. Mas-
3
todons were less tall but had relatively longer bodies.

Modern elephants live in hot places where they seem to have trouble
keeping cool, but m a n y m a m m o t h s lived in temperate or even cold
places. Cave paintings in France and Spain show m a m m o t h s w i t h long
hair, which they may have needed for w a r m t h during the Ice Ages.
Frozen carcases of Mammuthus primigenius have been found embed-
ded in ice in Siberia: the m a m m o t h s seem to have fallen into crevasses,
died and frozen, and to have remained frozen until they were found.
They have long black hair, just as shown in the cave paintings, and
they also have an 8-centimeter layer of fat under the skin. Both the
hair and the fat may have been useful as heat insulation. Modern el-
ephants can maintain their body temperatures in the w a r m climates
of Africa and India, and also in zoos in temperate countries, but m a m -
m o t h s inhabited m u c h colder e n v i r o n m e n t s and probably needed extra
insulation.
Most extinct m a m m a l s , including the m a m m o t h s , seem unspectac-
ular in comparison with dinosaurs. The saber-tooths [Smilodon], which
160 GIANT MAMMALS
preyed on m a m m o t h s , were impressive, but they were only about the

size of modern lions. The giant Irish deer [Megaloceros] was smaller in
the body than a moose though its antlers grew to a span of 3.5 meters.
The giant ground sloth [Megatherium) of South America was enor-
mously larger than any modern sloth but its estimated mass (3 tonnes)
is no more than that of the biggest modern rhinoceros.
There are various other extinct m a m m a l s of modern rhinoceros size,
and just one that is enormously larger. It is Indricotherium (formerly
called Baluchitherium), a hornless rhinoceros from Mongolia. It is hard
to be sure of its m a x i m u m size because no complete skeleton has been
found, only odd bones from specimens of various sizes. Only two neck
vertebrae and a foot bone (a metacarpal) seem to come from the biggest
size of skeleton. These three bones have been drawn to scale in figure
12.1, and the rest have been scaled up from smaller skeletons. The
sizes of some ribs and vertebrae have had to be guessed because no
specimens were found of those particular bones. T h u s the evidence for
the size of the animal in figure 12.1 is shaky: a small error of judgment
could have made it badly wrong. Since no better evidence is available,
I will assume that the figure is accurate. It shows an animal 5.3 meters
tall, m u c h taller than any elephant.
F I G U R E 12.1. A r e c o n s t r u c t i o n of t h e l a r g e s t Indricotherium. From Granger

a n d G r e g o r y (1935).
GIANT MAMMALS 161
The scientists responsible for the picture estimated the mass of the
animal to be 20 tonnes, but I think it m a y have been even more. T h e
head and body (excluding the tail) are 9.2 m e t e r s long, measured along
the curve of the back. T h e same m e a s u r e m e n t in 0.75-tonne African
buffalo is 2.6 meters. T h e Indricotherium has a body of roughly buf-
falo-like shape, so if it ws (9.2/2.6) t i m e s as long as the buffalo it was
(9.2/2.6) t i m e s as heavy: about 34 tonnes. I have tried calculating its
1
mass in other ways and obtained even larger estimates. If the resto-
ration is accurate, Indricotherium had about the same mass as Apa-
tosaurus (figure 1.7).
When we discussed the heat balance of large dinosaurs (chapter 7)
we were uncertain whether they had reptile-like or m a m m a l - l i k e m e -
tabolism. We concluded that a large sauropod w i t h m a m m a l - l i k e me-
tabolism would have to evaporate a lot of water to avoid overheating
in hot climates. Indricotherium is obviously a m a m m a l and presum-
ably had m a m m a l - l i k e metabolism, but the climate in Mongolia m u s t
have been reasonably cool in its time, as it is now. T h e continents had
by then reached their present positions on the earth's surface.
Indricotherium is the only land-living m a m m a l k n o w n to have grown
to the size of large sauropods, but there are bigger m a m m a l s in the sea.
The Blue whale [Balaeonoptera musculus) grows to an average adult
mass of about 100 tonnes and is the biggest animal k n o w n to have
lived, at any time. Its n u m b e r s have been very seriously reduced by
whaling but, happily, it survives, so it needs no further discussion in
this book on extinct giants.
Principal Sources
Savage and Long (1986) d e s c r i b e t h e e x t i n c t g r o u p s of m a m m a l s . E c o n o m o s (1981)

d i s c u s s e s Indricotherium. G r a n g e r a n d G r e g o r y (1936) is t h e s o u r c e for figure 1 2 . 1 .
E c o n o m o s , A. C. 1 9 8 1 . T h e largest l a n d m a m m a l , journal of Theoretical Biology

89:211-215.
G r a n g e r , W. and W. K. G r e g o r y . 1935. A r e v i s e d r e s t o r a t i o n of t h e s k e l e t o n of Balu-
chitherium, g i g a n t i c fossil r h i n o c e r o s of C e n t r a l A s i s . American Museum Novi-
tates 7 8 7 : 1 - 3 .
Savage, R. J. G. a n d M. R. Long. 1986. Mammal Evolution: An Illustrated Guide.
L o n d o n : British M u s e u m ( N a t u r a l H i s t o r y ) .
XIII
Epilogue
T HIS BOOK about gigantic animals has highlighted some of the spe-
cial problems of large size, m a n y of which depend on the rule of
squares and cubes. T h e weights of geometrically similar animals of dif-
ferent sizes are proportional to the cubes of their lengths, but the areas
of corresponding body surfaces are proportional only to the square of
length: an animal twice as long as another animal of the same shape
is eight times as heavy but has only four t i m e s the area. This is why
large animals get bogged down in m u d more easily than small ones
(chapter 3): their weights are proportional to the cube of length but the
areas of the soles of their feet only to the square. It is why large flying
animals m u s t fly fast, to keep themselves airborne, and may have trou-
ble taking off (chapter 8): their weights are proportional to the cube of
length but the areas of their wings only to the square. It is also why
large animals are not as athletic as small ones (chapter 4): the forces
that act on t h e m in dynamically similar activities are proportional to
body weight and so to the cube of length but the strengths of bones
and muscles (which depend on cross-sectional area) only to the square.
This last example is a little more complicated than the others because
the transverse force that a bone can stand, acting on its end, is not
directly proportional to area but to the ratio Z / x : however, Z / x is pro-
portional to the square of length in geometrically similar animals.
We depart further from the simple rule of squares and cubes in ques-
tions of heat balance (chapter 7). Rates of metabolic heat production
are not proportional to body mass but more nearly to (body m a s s ) . 07S
Rates of loss of heat, for equal temperature differences between body

and environment, are not simply proportional to surface area but de-
pend also on the thickness of skin and any additional insulating layer.
However, the effect of the relationships is that large animals can be
more effectively endothermic ("warm-blooded") than small ones. Shrews
EPILOGUE 1 63
and other small e n d o t h e r m s need relatively thick fur or feathers but

large ones such as elephants need no fur at all and very large endo-
therms might be liable to overheat. Similar conclusions apply to egg-
shells (chapter 11) because the m a t h e m a t i c s of gas diffusion resembles
the m a t h e m a t i c s of heat conduction: big eggs (such as those of os-
triches) need more porous shells t h a n small ones (such as those of spar-
rows) to allow gases to diffuse in and out fast enough to sustain their
metabolism. Finally, large ectothermic ("cold-blooded") animals take
longer than small ones to equilibrate to a new environmental temper-
atures. Small lizards approach equilibrium w i t h i n a few m i n u t e s but
a large ectothermic dinosaur would take m a n y days.
Those arguments depend on assumptions of geometric similarity, but
large animals are not geometrically similar to small ones. Elephants
are not the same shape as shrews, nor are albatrosses the same shape
as hummingbirds. Masses of geometrically similar animals would be
proportional to (bone circumference) and the masses of real m a m m a l s
1
are proportional to (bone c i r c u m f e r e n c e ) ' (figure 2.5). Wing loadings

27
of geometrically similar birds would be proportional to (body m a s s ) 0 3 3
and actual wing loadings are proportional to (body m a s s ) 0 4 0

both for
marine soarers and for land soarers (figure 8.8). Heating and cooling
time constants in water for geometrically similar animals made of the
same materials would be proportional to (body m a s s ) , and the ob-
067
served time constants for different sized reptiles are proportional to

(body m a s s ) and (body m a s s )
067 071
(figure 7.4). T h e deviations from geo-
metric similarity modify the effect of differences of size but do not
cancel the general trends.
These points form part of the message of this book, but there is an-
other very important part: engineering theory, designed for application
to man-made structures, can also help us to understand the structure
and behavior of animals.
I used an idea that had its origin in shipbuilding to define equivalent
running speeds for animals of different sizes, and information from soil
mechanics to assess the danger of dinosaurs getting bogged down in
mud or sand (chapter 3). I used m e t h o d s developed by engineers to cal-
culate stresses in structures such as bridges w h e n I discussed h o w ath-
letic dinosaurs could have been (chapter 4), and again in calculations
about dinosaur neck ligaments (chapter 5). T h e calculation of forces
for a collision of dome-headed dinosaurs could have been applied to
automobiles, and the discussion of Parasaurolophus' voice used simple
acoustics (chapter 6). T h e discussion of dinosaur heat balance (chapter
7) used theory such as heat engineers use w h e n designing central heat-
ing systems. I used aerodynamics developed for application to aircraft
to show how pterosaurs probably flew (chapter 8) and similar theory
164 EPILOGUE
applied to water instead of air to show how ichthyosaurs and plesio-

saurs m a y have s w u m (chapter 9). Geophysical information was needed
in the discussion of dinosaur extinction (chapter 10) and diffusion the-
ory in the discussion of birds' eggs (chapter 11).
Physics and engineering are as useful in the study of living animals
and of the h u m a n body, as in the study of dinosaurs and other extinct
animals. Physics is the basic science of m a t t e r and energy, and engi-
neering is physics applied to structures and machines. They and chem-
istry are the sciences that biologists need to explain the structure and
m e c h a n i s m s of living things.
INDEX
(italics indicate figures or tables)

A l b a t r o s s , 111, 114, 151 C e n t r e of g r a v i t y , 6 3 , 64, 65. 69
Allosaurus, 8, 10. 14 C e r a t o p i a n s , 13
Alvarez, D r s . L a n d W., 141 C h e e k s , 12
Amphibolurus. 104 C h e w i n g , 12
Anapsida, 7 C l a s s i f i c a t i o n , 7, 8, 9
Anatosaurus, 12, 13. 15, 82 C o l b e r t , Dr. E., 18
A n d e r s o n , Dr. ]. F., 20 Cold-blooded, 90
A n d e r s s o n , Dr. M., 83 C o m e t s , 148
Angle of a t t a c k , 109 Compliance, 86
A n k y l o s a u r s , 14 Compsognathus, 1, 3, 6, 8, 10. 14
A n t l e r s , 74 C o n d o r , 111, 151
Apalosaurus. 9, 10. 14, 50, 6 2 ; foot of, Cooling, 95
30. 3 4 ; n e c k of, 6 6 ; tail of, 70 Copulation, 57
Archaeopteryx, 150 Corythosaurus, 82, 83
Archimedes' Principle, 19 C o t t , Dr. H., 19
Archosauria, 7 C r e s t s , 82, 120
Argentavis, 151 C r e t a c e o u s Period, 15, 141
A s p e c t ratio, 111, 111. 126 C r o c o d i l e , 10. 19
A s t e r o i d s , 143 C r u m p l e zones, 82
Cryptoclidus, 130
Bakker, Dr. R., 62, 93 C y c a d , 14
Baluchitherium, 160
Bending m o m e n t s , 47 D e e r , 74, 75; g i a n t , 160
Bends, t h e , 129 Deinonychus, 69
Birds, giant, 150 D e n s i t y , 19, 127
Blood p r e s s u r e , 61 Diatryma. 152
Bones, d i m e n s i o n s of, 20; t u b u l a r 113 Dimorphodon, 108
Brachiosaurus. 1, 2, 6, 11, 19, 22; Dinornis, 152
c e n t r e of gravity, 6 3 , 6 5 ; h e a t Diplodocus, 9, 11. 62, 6 5 n e c k of, 60,
;
b a l a n c e , 99, 104 n e c k , 60
; 66, 6 7 ; tail of, 70
B r a m w e l l , Dr. C , 120 D i s p l a y , 77
Brontosaurus, 9 D i v i n g , 127
B u o y a n c y , 60, 127 D o m e - h e a d e d d i n o s a u r s , 14, 80
166 INDEX
Drag, 109 Indricotherium, 160

Dryoplosaurus, 71 I n d i u m , 142. 146
D u c k - b i l l e d d i n o s a u r s , 4, 82
Jurassic period, 14
Ectotherms, 90
Eggshells, diffusion t h r o u g h , 156; of K a n g a r o o s , 69
d i n o s a u r s , 4, 100; of e l e p h a n t birds, Kori b u s t a r d , 111, 151
156 Kronosaurus, 129
Elasmosaurus, 129
E l a s t i n , 64 Lambeosaurus, 83
E l e p h a n t , 18, 34, 102 Lift, 109
E l e p h a n t birds, 152, 156 L i g a m e n t s of n e c k , 64, 66, 67
E n d o t h e r m s , 90, 106, 126 Loch N e s s m o n s t e r , 138
Euoplocephalus. 71 Logarithms, 23
E x t i n c t i o n , 141 Lungs, 5 3
Fighting, 73 M a l o i y , Prof. G. M. O., 21

F i t t e s t , s u r v i v a l of, 73 M a m m o t h s , 159
Flight, 106; u n d e r w a t e r , 131 M a r i n e r e p t i l e s , 122
Flippers, 125, 130 M a s s e s of d i n o s a u r s , 16, 25; of g i a n t
F o o t p r i n t s of d i n o s a u r s , 10, 27, 28, 30, birds, 153; of Indricotherium, 161; of
31; of m o a s , 155; of m o d e r n a n i m a l s , m o d e r n a n i m a l s , 18, 122, 159, 161
32, 35; s u p p o s e d , of p t e r o s a u r s , 107 M a s t o d o n s , 159
Forces on feet, 44, 4 7 , 53 Megaloceros, 160
Fossils, 2, 5 Megatherium. 160
F r e q u e n c y of l i m b m o v e m e n t s , 136; M e t a b o l i c rate, 9 1 , 92, 100, 101. 162
resonant, 86 M e t e o r i t e , 141
Frigate birds, 115, 120 M o a s , 152
Froude n u m b e r , 43 M o m e n t u m , angular, 69; linear, 118
Fur, 94, 106 M o s a s a u r s , 128
Mud, 30
Gaits, 39
Genes, 73 N a m e s of species, 6
Genus, 6 N a s a l passages, 8 5
Giraffe, 1, 2, 62 N e c k s of d i n o s a u r s , 6 0 ; of p l e s i o s a u r s ,
G l i d e r s , 109 137
G r a v i t a t i o n a l a c c e l e r a t i o n , 17 N e w t o n ( u n i t of force), 17
Hadrosaurs, 82 Oldman Formation, 94

H a l l - M a r t i n , A., 20 O s t r i c h , 152
H a l s t e a d , Dr. B., 58 O r n a m e n t s , 83, 120
H a t c h i n g , 157 O r n i t h i s c h i a n , 10, 11, 12
Heat capacity, 96 O r n i t h o p o d , 12, 28, 68
H e r d s of d i n o s a u r s , 42 O z o n e , 146
Horned dinosaurs, 13
H o r n s , 74, 78 P a c h y c e p h a l o s a u r s , 14, 80, 81
Hydrofoil, 125 Pachyornis, 153, 154
Hypsilophodon, 81 P a d i a n , Dr. K., 106
Parasaurolophus, 83, 86
I c h t h y o s a u r s , 122, 123, 126 Pelvic girdle, 12
Iguanodon, 12, 13. 33. 6 8 ; foot of, 30, P e n g u i n s , 131, 135, 137
3 4 ; h e a t b a l a n c e , 9 9 ; s k u l l , 10 P l e s i o s a u r s , 129. 130. 132. 133, 134
INDEX 1
P o p u l a t i o n , 105 Strain, 45
Porpoising, 126 S t r e a m l i n i n g , 124
P r e d a t o r s and prey, 93 Stress, 46, 5 1 , 52
Protoceratops, 4, 77 S t r e n g t h , 44, 153
Psittacosaurus, 1, 3, 4, 5, 6, 12, 13, 100 S t r i d e l e n g t h , 34, 36, 38
Pteranodon, 107, 108, 113, 119, 120, Styracosaurus, 79, 80
151 Supersaurus, 1
P t e r o s a u r s , 106
Tails, 67
Quetzelcoatlus, 107, 151 Take-off, 111
T e e t h , 8, 12, 13
Radioactive dating, 5
T e m p e r a t u r e , body, 90, 104, 126
Rain, acid, 146
Tendons, 68
R a t i t e s , 152
Teratornis, 151
Reptile classification, 7, 8
Thaumatosaurus, 130
R e s o n a n c e , 86
Thecodonts, 7
Rhamphorhynchus, 107
T h e r m a l s , 114, 116
Running, 39
T h e r o p o d s , 8, 28
Russell, Dr. D., 20
T h u l b o r n , Dr. R., 42
T i m e , geological, 7
Sabre t o o t h , 159
T i m e c o n s t a n t , 96, 98, 101
Safety factor, 153
T o n n e , 17
S a u r i s c h i a n , 10. 11, 12
T r a d e w i n d s , 115, 119
Sauropods, 9, 29
Triceratops, 13, 14, 15, 57, 6 4 ; h o r n s
Sea lions, 131
of, 75, 80
Section m o d u l u s , 47, 76
T u n a , 124
Sexual s e l e c t i o n , 8 3
T u r t l e s , 131, 137
Shark, 124
Tyrannosaurus, 8, 11, 12, 14, 15, 30,
Sheep, 75, 76
3 4 , 49
Similarity, d y n a m i c , 36, 5 0 ; g e o m e t r i c ,
163
Ultrasaurus, 1
Skin i m p r e s s i o n s , 4, 94
Sloth, giant, 160
Smilodon, 159 Voice, 88
Soaring, 113 V o l c a n o e s , 144, 146
Sound production, 85 V u l t u r e s , 115
Species, 6
Speed, d i m e n s i o n l e s s , 3 5 ; of flight, W a d e , Dr. M., 42
112; of r u n n i n g , 3 3 , 40, 4 1 , 9 1 ; of Walking, 39
s w i m m i n g , 123, 134, 137 Warm-blooded, 90
S q u a r e s and c u b e s , 162 W a t e r loss, 102
Stegoceras, 76 W e a p o n s , 70, 74, 80
Stegosaurus. 14, 6 3 , 7 1 , 103 Whitfield, Dr. G. R., 120
S t o m a c h c o n t e n t s of d i n o s a u r s , 4, 8, 9; Widowbirds, 83
of o t h e r a n i m a l s , 119, 123, 153; W i n g l o a d i n g , 111, 115, 117
stones, 9 W i n g s , 106

R. McNeill Alexander-Dynamics of Dinosaurs and Other Extinct Giants-Columbia University Press (1989) PDF

Uploaded by

Copyright:

Available Formats

R. McNeill Alexander-Dynamics of Dinosaurs and Other Extinct Giants-Columbia University Press (1989) PDF

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

R. McNeill Alexander-Dynamics of Dinosaurs and Other Extinct Giants-Columbia University Press (1989) PDF

Uploaded by

Copyright:

Available Formats

DYNAMICS OF

Columbia University Press

I. Introducing the Dinosaurs 1

XII. Giant M a m m a l s 159

xm. Epilogue 162

T H I S is a book about big, extinct animals. Most of it is about di-

Introducing the Dinosaurs

formed and introduces some of the best-known dinosaurs. It is an

stayed together. Many other dinosaur finds have been of scattered or

F I G U R E 1.1. Brachiosaurus w i t h a h o u s e a n d a n a d u l t giraffe d r a w n t o t h e

FIGURE 1.2. J u v e n i l e Compsognathus ( b e h i n d ) a n d Psittacosaurus (right), w i t h

sand becomes sandstone, m u d becomes shale and calcium carbonate

F I G U R E 1.3. A Psittacosaurus s k e l e t o n a s i t w a s f o u n d , with enough rock

already buried, so the small one finished up in a higher layer of rock.

want to write about Brachiosaurus brancai (the best-known species)

TABLE 1.1. T h e divisions of geological t i m e .

Mesozoic Cretaceous 140

Palaeozoic (six p e r i o d s ) 570

N O T E : D i n o s a u r s lived during the M e s o z o i c era.

appearance, making particular m e n t i o n of their teeth and other evi-

TABLE 1.2. Classification of dinosaurs mentioned in this book and

TABLE 1.3. How reptiles are classified.

F I G U R E 1.6. S k u l l s of (a) a c r o c o d i l e ; (b) a s a u r i s c h i a n d i n o s a u r [Compsog-

culiarity of ornithischians is an extra bone (the predentary) at the front

FIGURE 1.10. P e l v i c g i r d l e s o f a s a u r i s c h i a n {Tyrannosaurus) a n d a n o r n i t h i s -

FIGURE 1.13. Triceratops ( c e n t e r ) a n d Stegosaurus (right), d r a w n f r o m m o d e l s

years later, late in the Cretaceous, Tyrannosaurus, Anatosaurus, and

Bakker, R. T. 1986. The Dinosaur Heresies. N e w Y o r k : M o r r o w .

of dinosaurs in chapter 1 were based on fossil bones,

.L fitted together to form complete skeletons. These skeletons tell

hindered by air resistance or anything else, increases by 10 meters per

TABLE 2.1. Masses (in tonnes) of some modern animals.

Blue whale, Balaenoptera musculus 91 110

African elephant, Loxodonta africana 5.45 2.77

Hippopotamus, Hippopotamus amphibius 2.52 2.13

Black rhinoceros, Diceros bicornis 1.17 1.08

Eland, Taurotragus oryx 0.84 —

African buffalo, Syncerus caffer 0.75 —

L i o n , Panther a leo 0.18 0.15

Human, Homo sapiens 0.07 0.05

S O U R C E S : w h a l e : N i s h i w a k i 1950; e l e p h a n t : Laws 1966; others: M e i n e r t z h a g e n 1938.

Colbert's method was difficult to perform accurately because it de-

FIGURE 2.1. A p p a r a t u s for m e a s u r i n g t h e v o l u m e s o f m o d e l d i n o s a u r s .

like m a m m a l s than like crocodiles, lizards, and tortoises. T h e Ander-

F I G U R E 2.2. (a) a 3 . 9 - k i l o g r a m S e c r e t a r y b i r d [Sagittarius serpentarius a n d (b)

FIGURE 2.3. The skeleton o f Brachiosaurus brancai, showing the n a m e s of

to errors due to different quadrupeds supporting different fractions of

distances along both scales are proportional to the logarithms of the

body mass in kg = 0.000084 (total of circumferences in m m ) 2 73

the circumferences of their bones. However, appearances can be de-

body mass in kg = 0.00016 (femur circumference in mm) ™. 2

This underestimates the masses of kangaroos and overestimates the

TABLE 2.2. Masses (in tonnes) of dinosaurs.

(1962) (1985) (1985)

N O T E : T h e Colbert e s t i m a t e s , a s originally published, w e r e based o n a n a s s u m e d d e n s i t y o f 9 0 0

It is hard to decide w h i c h of the mass estimates to prefer, in table

A l e x a n d e r , R. M c N . 1985. M e c h a n i c s of p o s t u r e a n d gait of s o m e large d i n o s a u r s .

O F A L L that remains of dinosaurs, their footprints bring the animals

F I G U R E 3 . 1 . D i n o s a u r footprints found at W i n t o n , Q u e e n s l a n d , from T h u l -