angiosperms possessed adaptations that were new and potentially
revolutionary. But they needed some edge—an opening for
breaking out of the confines imposed by the older groups to start
proliferating species. Herbivorous dinosaurs gave them that initial
break.
Flowering plants and low-feeding beaked dinosaurs must have
co-evolved in a mutually beneficial way. As more and more new
kinds of Cretaceous beaked dinosaurs entered the system, more
and more angiosperm families evolved. From the meek beginning
of a few Early Cretaceous species, the angiosperms grew into a
mighty clan by the Late Cretaceous, boasting more species than
conifers and cycadeoids combined. When the Cretaceous ended,
massive extinctions again swept through the terrestrial habitats.
Cycadeoids disappeared, but angiosperms not only survived, they
increased their share of the species count in epoch after epoch down
to the present day.
Modern angiosperms no longer depend upon intense low
feeding for their advantage over conifers and ferns. They now
contain hundreds of specializations for every habitat from tall cli-
max forests to swamps and deserts, windswept mountain meadows
and bare rock faces. But in those first critical years of their evo-
lutionary history, the angiosperms were struggling newcomers.
Square snouts and pincerlike beaks helped the flowering plants beat
the floral competition and establish the angiosperms as the fastest-
evolving plant group. There was of course no plan in this. Stego-
WHEN DINOSAURS INVENTED FLOWERS I 197
saurus didn't die out purposely to permit its low-cropping cousins
to take over and make the world fit for flowers. Wide-mouthed
ankylosaurs didn't plan to munch down the competition. It was all
serendipitous. Nonetheless, because of the way they suffered ex-
tinction and then rebuilt their herbivorous groups, the dinosaurs
played a central role in one of the grandest dramas of the flora. In
their way, dinosaurs invented flowers. Without them, perhaps our
modern world would yet be as dull green and monotonous as was
the Jurassic flora.
198 I THE HABITAT OF THE DINOSAURS
PART 3
DEFENSE,
LOCOMOTION,
AND THE CASE
FOR WARM-BLOODED
DINOSAURS
10
THE TEUTONIC
DIPLODOCUS: A LESSON IN
GAIT AND CARRIAGE
nyone who doesn't believe that God was looking out for
A
America at the turn of the century should look at the dates
of key American victories in war and science. On July 4, 1898,
Admiral Sampson announced a "Birthday present for the Nation,"
the complete victory of the fleet over the Spanish squadron in Cuba.
(Commodore Schley actually won the battle. Sampson was away
conferring with generals—but claimed the credit.) On July 4, 1899,
Arthur Coggeshall found Diplodocus carnegiei. On July 4, 1900, El-
mer S. Riggs, hunting dinosaurs for Marshall Field's Chicago mu-
seum, found the first-known Brachiosaurus, king of the brontosaurs,
a giant that dwarfed even Brontosaurus excelsus.
American museums were erecting dinosaur skeletons as fast
as American shipyards erected new steel battleships to protect the
fledgling star-spangled empire. Europe viewed both developments
with mixed admiration and alarm. For a century Old World sci-
entists had been digging and studying dinosaurs, but no one had
found Jurassic giants nearly as complete as the ones that tumbled
out of almost two dozen American quarries, starting in 1878. When
the first Diplodocus, named for the American millionaire Andrew
Carnegie who funded its discovery, arose on its metal scaffolding
in Pittsburgh, John Bell Hatcher, in charge of the operation, di-
rected the placement of the thigh bone into the hip socket very
carefully. He drew upon the anatomical expertise of Marsh, Cope,
THE TEUTONIC DIPLODOCUS: A LESSON IN GAIT AND CARRIAGE
201
Andrew Carnegie's ninety-foot-long
Diplodocus from Sheep Creek, Wyoming
and other Americans who had engaged in two decades of intense
study. Hatcher decreed a vertical stride, with the thigh bone
swinging fore and aft directly under the hips, for Diplodocus car-
negiei. And the other American museums consequently agreed.
Riggs's Brontosaurus went up that way in Chicago, as did Osborn's
in New York. These Americans were convinced that the great di-
nosaurs strode through their Mesozoic world with the upright gait
and carriage that are characteristic of the biggest African elephants
today.
America, which had had to import all its scientific apparatus,
and had sent its scholars to England and Germany for doctorates
only decades earlier, now began to export scientific wealth.
On one of his frequent visits in British high society, Andrew
Carnegie met with Edward, Prince of Wales, the future Edward
VII. Aware of Carnegie's enthusiasm for the exploration of dino-
saur sites in Wyoming as an aspect of his new-found passion for
public service, the Prince of Wales suggested Carnegie might be
pleased to have his people find another Diplodocus for the British
Museum, which had no complete specimen.
Back in America, William J. Holland, director of the Pitts-
burgh Museum, was aghast at Carnegie's request. New quarries not
already being worked by other new American museums were ex-
tremely difficult to find, and it would be impossible to guarantee
quick delivery. Holland proposed a complete plaster replica in-
stead. So, in due course, the Pittsburgh technicians assembled
beautifully accurate plaster casts of Diplodocus carnegiei and shipped
them to London complete with instructions for assembly. A char-
acteristic American approach—the prefabricated, instant dinosaur
kit. Soon Carnegie was besieged by envoys from Berlin, Vienna,
and St. Petersburg for matching gifts of a Diplodocus. And Andrew
was delighted to comply. Within a few years nearly every major
European capital had its own prefab Diplodocus.
Once assembled, most of these European Diplodocus replicas
were posed in mid-stride, with the same high-hipped posture pre-
scribed by Hatcher and the other American experts. This recon-
struction bore important implications for dinosaur biology, precisely
because no living species of reptile walked that way. Most pres-
ent-day lizards scuttle over the ground with their thighs sticking
out sideways. Their hind limb strokes back and forth in horizontal
THE TEUTONIC DIPLODOCUS: A LESSON IN GAIT AND CARRIAGE I 203
arcs, parallel to the ground. Hence, these modern lacertilians are
decidedly low-slung in posture, with a ground-hugging configura-
tion that allows them only slight clearance. Crocodilians and cha-
meleon lizards are a bit more upright in carriage, raising their bellies
higher off the trackway. Yet their elbows and knees still stick out
sideways more than those of an elephant, rhino, or most other large
mammals. Endowing the Diplodocus with elephant-style posture was
therefore a clear statement that the biomechanics of dinosaurs were
unlike those of any living reptile. Diplodocus's posture as envi-
sioned by Hatcher, Osborn, and Riggs was equal to the most ad-
vanced mammalian adaptive machinery.
Enter the Germans. No culture had a more illustrious nine-
teenth-century tradition of paleontological scholarship. A Ger-
man, Hermann von Meyer, had first recognized the unity of all
the great Mesozoic creatures we now call dinosaurs. And German
anatomists were acknowledged worldwide as the best in labora-
tory dissections and microscopy. In the early 1900s, Germany was
a new and ambitious nation, and it was perhaps to be expected
that a certain chauvinism should manifest itself in many different
areas, including the scientific. It was not surprising that German
paleontologists didn't immediately accept the conclusions about the
posture of dinosaurs advocated by the Americans. What was sur-
prising was the condescending tone the Germans resorted to when
they published their scathing criticism of Carnegie's Diplodocus. The
Germans insisted that the Americans had missed the point when
they put elephant's legs on a dinosaur. Diplodocus was a genuine
reptile, the elephant a genuine mammal, and nature did not mix
the two. Tornier, the dinosaur expert in Berlin, described his ver-
sion of the corrected Diplodocus: it was portrayed in a slinking pose,
with the thigh and arm sticking out sideways, and the belly close
to the ground. The Berlin school proclaimed this was a proper
reptilian posture for a proper reptilian body.
The Americans did not concur. They had a well-earned na-
tional reputation for a hard-headed approach to the functions of
machinery. Hatcher had handled scores of fossil hip joints. He knew
that Diplodocus's thigh had a cylindrical surface at the joint that faced
predominantly upward and forward. Diplodocus's hip bone con-
tained a deep socket at the joint, whose surface correspondingly
faced mostly downward and backward. Put the thigh bone into the
204 | DEFENSE, LOCOMOTION, AND THE CASE FOR WARM-BLOODED DINOSAURS
Diplodocus in a rut. Carnegie
Museum's Professor
Holland poked fun at the
German reconstruction of
Diplodocus with its
outspread knees. Holland
remarked that the poor
German Diplodocus would
have had to find giant ruts
to run in because its rib
cage was so deep. (The
diagram shows a cross-
section view at the hip
joint.)
How Carnegie's men
mounted Diplodocus. The
Americans put together the
hind limb the right way—
with the thigh bone in a tall,
erect posture.
hip socket, and only one correct fit was possible: the hind leg stood
vertically, with the knee facing directly forward. The Diplodocus's
knee did not sprawl sideways like that of a "genuine lizard."
American scholars, one after another, rebutted the arguments
of the Germans. A key point all too often forgotten today was made
in the course of this debate. Dinosaur biology cannot be recon-
structed by assuming these beasts were merely "good reptiles."
By 1920, the Great Trans-Atlantic War of the posture of Di-
plodocus was over. The Americans clearly had the better of it. Ex-
peditions from Berlin had found fabulous Brachiosaurus graveyards
THE TEUTONIC DIPLODOCUS: A LESSON IN GAIT AND CARRIAGE | 205
in German East Africa before World War I, and as the postwar
research was published, the German scholars' restoration of Brach-
iosaurus became like Carnegie's, with a nearly vertical, elephant-
like posture. Incontrovertible evidence for the correctness of the
Pittsburgh-style hip joint came during the 1930s and 1940s, when
brontosaur footprints were found in abundance impressed into the
Cretaceous limestone of Texas. The right and left hind prints proved
to have been made very close to the trackway centerline, and
therefore without question the two thighs had swung in great ver-
tical arcs close to one another under the animal, exactly as Hatcher
had reconstructed them. Indeed, all of the dinosaur fossil track-
ways without exception showed the very same narrow-tracked gait,
evidencing no splaying out of the knees. No dinosaur splayed its
knees or toed out its feet. Duckbill dinosaurs actually toed in their
hind feet, like enormous pigeons.
Sad to relate, some modern reconstructions done in the 1960s
and 1970s still portray dinosaurs everting their knees and planting
their hind feet down, wide-set, with right and left hind paws spread
far out to the side—like enormous lizards. An entire series of
postcards in the British Museum reproduces paintings with such
essentially dislocated dinosaur hips and ankles. Andrew Carnegie
and Prince Edward knew better in 1906.
As a student at Yale in the sixties, I observed that nearly
and narrow trackway. But the forelimbs were a different matter
entirely. In the Great Hall of the Peabody Museum, trailing be-
hind the nobel strides of Marsh's Brontosaurus excelsus, stood a finely
preserved horned dinosaur, Centrosaurus. Professor Richard Swann
Lull had mounted it in 1929- He had been with Osborn at Como
and was highly regarded. He gave the centrosaur tall, erect hind
limbs, but the elbows were mounted lizard-style, sticking out
sideways, and the upper arms paralleled the ground. I marveled at
this curious combination, which looked like two totally different
locomotor apparatuses welded together at mid-torso. Further-
more, Lull had also published a monograph on the horned dino-
saur in which he employed this mismatched front end as an
argument for a slow and plodding gait in these animals. But Marsh
had already seen it otherwise as early as 1896. He had published
drawings of horned dinosaurs with fully erect carriage in both fore-
206 | DEFENSE, LOCOMOTION, AND THE CASE FOR WARM-BLOODED DINOSAURS
quarters and hind. Marsh's Triceratops imparted a light-footed air
to the huge three-horned beast, as though it were about to go
trotting off the page, head down, to charge its mortal enemy, Ty-
rannosaurus. Lull completely reversed Marsh's ideas about these
forelimbs.
But why had views on the forelimbs of dinosaurs changed? I
could find no good reason. Smithsonian scientists endowed their
Triceratops, a skeleton excavated by Hatcher, with bowed-out el-
bows because, they said, the elbow's huge "funny bone" would be
of use only in a sprawling posture. This made no sense to me. The
"funny bone," properly called the olecranon, is a projection on the
lower arm bone (ulna) to provide the elbow-opening muscles le-
verage for their work. A large olecranon implies the elbow joint
can be opened with great force. Now, turtles, crocs, and lizards
pos
sess splayed-out elbows, but they all have short olecranons. The
rhino's forelimbs stride vertically, and its olecranon is big. Tricer-
atops possessed a large, rhino-style funny bone. Why wasn't Tri-
ceratops accorded a rhino-style posture?
Al Romer, the greatest dinosaur anatomist alive in the 1960s,
explained the anomaly by arguing that evolution had worked faster
on the hind legs than the fore. All primitive dinosaurs, as Romer
told the story, had been two-legged bipeds, standing on hind limbs
only. Thus when some families later dropped back down onto all
fours, they didn't bother to rearrange their elbows to match their
knees. This account was in all the textbooks, but I could not ac-
cept it. Yale had a program—the Scholar of the House—in which
an undergraduate could dedicate one full year exclusively to re-
search. For my project, I focused on the problem of the alignment
of the forelimbs of dinosaurs.
It was a shoestring operation. My parents gave me a tiny
handwound movie camera, and with Yale funds I bought the alli-
gators and lizards that I kept in the museum basement. I'd lie on
my stomach watching a lizard or 'gator walk around on an old rug,
and built up a library of motion-analysis film of a dozen species.
These filmings had their dangers. No three-foot 'gator was afraid
of a Yale undergraduate, and I had to remember to move smartly
when an open mouth filled the viewfinder. Usually I was in time
to avoid having my nose bitten.
Alligators are dinosaur uncles—relatives of the direct ances-
THE TEUTONIC DIPLODOCUS: A LESSON IN GAIT AND CARRIAGE | 207
The riddle of the mismatched legs. Living species have a range of postures—
most lizards sprawl, crocodilians have a more upright, semierect stance, and
most big mammals have a fully erect carriage. But orthodox dinosaurs—like
Yale's Centrosaurus—had front ends that didn't align with the rear ends.
tors of early dinosaurs—and as such they should be living repre-
sentatives of the ancestral dinosaurs' forelimb arrangement. I was
therefore surprised at the upright arrangement of the alligators'
forelimbs. They kept their elbows close to their sides without
spreading nearly as much as most lizards do. So I labeled this pos-
ture "semi-erect," to set it apart from the "fully erect" posture of