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The
PRINCETON
FIELD GUIDE to
DINOSAURS
The
PRINCETON
FIELD GUIDE to
DINOSAURS
3RD EDITION
GREGORY S. PAUL
press.princeton.edu
ISBN 978-0-691-23157-0
ISBN (e-book) 978-0-691-23156-3
10 9 8 7 6 5 4 3 2 1
CONTENTS
Preface6 Growth56
Acknowledgments6 Energetics57
Gigantism61
Introduction9 Mesozoic Oxygen 64
History of Discovery and Research 9 The Evolution—and Loss—of Avian
What Is a Dinosaur? 14 Flight64
Dating Dinosaurs 15 Dinosaur Safari 66
The Evolution of Dinosaurs and If Dinosaurs Had Survived 67
Their World 16 Dinosaur Conservation 68
Extinction26 Where Dinosaurs Are Found 69
After the Age of Dinosaurs 28 Using the Group and Species Descriptions 76
Biology28
General Anatomy and Locomotion 28
Group and Species Accounts 83
Skin, Feathers, and Color 38
Dinosaurs83
Respiration and Circulation 40
Digestive Tracts 42 Basal Dinosaurs 83
Senses45 Theropods86
Vocalization46 Sauropodomorphs202
Genetics46
Ornithischians261
Diseases and Pathologies 46
Behavior47
Additional Reading 374
Brains, Nerves, and Intelligence 47
Index to Dinosaur Taxa 375
Social Activities 48
Reproduction49 Index to Formations 382
PREFACE
If I were, at about age twenty as a budding paleozoologist Cerro del Pueblo, Hanson, Jinhua, Clearwater, Tiaojis-
and paleoartist, handed a copy of this book by a myste- han, Tsagayan, Tropic Shale, Mackunda, Blackleaf, Pari
rious time traveler, I would have been shocked as well Aike, Ukureyskaya, Villar del Arzobispo, Tegana, Kal-
as delighted. The pages would reveal a world of new di- lamedu, Wapiti, Bungil, Demopolis, Portezuelo, Prince
nosaurs and ideas that I barely had a hint of or had no Creek, Ulansuhai, Ulaanoosh, Sânpetru, Shinekhudag,
idea existed at all. My head would spin at the revelation Kitadani, Zhanghe, Woodbine, Calizas de la Huérguina,
of the therizinosaurs such as the wacky feathered Beipi- Chaochuan, Xert, Maevarano. The sheer number of new
aosaurus and at the biplane flying dromaeosaurids, not dinosaurs would demonstrate that an explosion in dino-
to mention the little halszkaraptors with their duck-like saur discoveries and research, far beyond anything that
beaks, or the oversized shoulder spines of Gigantspinosau- had previously occurred, and often based on new high
rus, the neck spines of Amargasaurus, the brow horns and technologies, marked the end of the twentieth century
atrophied arms of bulldog-faced Carnotaurus, the furry going into the twenty-first.
adornments of Tianyulong and Kulindadromeus, the bris- Confirmed would be the paradigm shift already under
tly tail of Psittacosaurus, the extraordinary preservation way in the late 1960s and especially the 1970s that ob-
of the armor of Borealopelta, the bat-like membranous served that dinosaurs were not so much reptiles as they
wings of scansoriopterygids, the long-necked stegosaur were near birds that often paralleled mammals in form and
Miragaia, and the often psychedelic frilled horns of the function. Dinosaurs were still widely seen as living in tropi-
new stable of centrosaurine and chasmosaurine ceratop- cal swamps, but we now know that some lived through
sids. Even Triceratops has proven to have strange skin. polar winters so dark and bitterly cold that low-energy rep-
How about colossal new sauropods that gave the giant tiles could not survive. Imagine a small dinosaur shaking
baleen whales a run for their money in terms of sheer the snow off its hairy body insulation while the flakes melt
bulk and could feed over six stories high in tree crowns? It on the scaly skin of a nearby titanic dinosaur whose body,
is a particular pleasure to restore the skeleton of the once oxygenated by a birdlike respiratory complex and pow-
mysterious Deinocheirus, long known from only its colos- ered by a high-pressure four-chambered heart, produces
sal arms—its peculiar skeleton does not disappoint. And the heat needed to prevent frostbite.
who would have imagined it would become possible to In just the eight years since the appearance of the sec-
figure out the colors of some feathered dinosaurs? I would ond edition of this book, the number of dinosaur species
note the new names for some old dinosaurs, including my named has expanded about 15 percent relative to some
favorite, Giraffatitan. And that old Brontosaurus is back! 200 years of research—if anything, there was more updat-
And now there are three royal species of Tyrannosaurus: ing between this edition and the second than between that
T. rex, T. regina, and T. imperator. There would be the di- and the first. Producing this third edition has been satisfy-
nosaur-bearing beds with the familiar yet often exotic ing in that it has given me yet more reason to more fully
names: Tendaguru, Morrison, Chinle, Wessex, Wealden, achieve a long-term goal, to illustrate the skeletons of al-
Lameta, Oldman, Kota, Kirtland, Arundel, Djadokhta, most all dinosaur species for which sufficiently complete
Nemegt, Cloverly, Navajo Sandstone, North Horn, Hell material is available. These have been used to construct
Creek, Portland, Lufeng, Arundel, New Egypt, Lance. the most extensive library of side-view life studies of di-
Plus there are the novel formations, at least to my eyes nosaurs in print to date. The result is a work that covers
and ears: Yixian-Jiufotang, Tiourarén, Dinosaur Park, what is now two centuries of scientific investigation into
Eumeralla, Longjiang, Two Medicine, Anacleto, Painten, the group of animals that ruled the continents for over 150
Shishugou, Huincul, Aguja, Ischigualasto, Qingshan, million years. Enjoy the travel back in time.
ACKNOWLEDGMENTS
A complaint earlier this century on the online Dinosaur Brett-Surman, Philip Currie, John McIntosh, Robert Bak-
List by Ian Paulsen about the absence of a high-quality ker, Peter Galton, James Kirkland, Scott Persons, Rinchen
dinosaur field guide led to the production of the first and Barsbold, John Horner, Xu Xing, Frank Boothman, Dan-
then the second edition of this dinosaur guide. The excep- iel Chure, David Burnham, Kristina Curry Rogers, Alex
tional success of those editions combined with the continu- Downs, Steven and Sylvia Czerkas, Peter Dodson, David
ous flux of new discoveries and research led to production Evans, James Farlow, Jay Van Raalte, Hermann Jaeger,
of a third. Many thanks to those who have provided the Frances James, Tracy Ford, Catherine Forster, Rodolfo
assistance over the years that has made this book possible, Coria, John Foster, Mike Fredericks, Roland Gangloff,
including Kenneth Carpenter, Asier Larramendi, Michael Donald Glut, Mark Hallett, Jerry Harris, Thomas Holtz,
6
Acknowledgments
Peter Larson, Nicholas Hotton, Guy Leahy, Nicholas Lon- John Jackson, Michael Triebold, David Varricchio, Mat-
grich, James Madsen, Jordan Mallon, Charles Martin, Hal- thew Wedel, David Weishampel, Jeffrey Wilson, Lawrence
szka Osmólska, Teresa Maryanska, Octavio Mateus, Carl Witmer, Ben Creisler, Dan Varner, Jens Lallensack, Saswati
Mehling, Ralph Molnar, Markus Moser, Darren Naish, Bandyopadhyay, Ji Shuan, Clint Boyd, Mickey Mortimer,
Mark Norell, Fernando Novas, Kevin Padian, Armand Tyler Greenfield, Mikko Haaramo, John Schneiderman,
Ricqles, Timothy Rowe, Dale Russell, Scott Sampson, John and many others. I would also like to thank all those who
Scannella, Mary Schweitzer, Masahiro Tanimoto, Michael worked on this book for Princeton University Press: Rob-
Taylor, Matthew Lamanna, Boris Sorkin, Robert Telleria, ert Kirk, Kathleen Cioffi, Laurel Anderton.
Iguanodon
7
The feathery dinosaur Avimimus
INTRODUCTION
9
History of Discovery and Research
also headed northeast of Beijing, they might have made even knowledge of the beginnings of predatory dinosaurs. Also
more fantastic discoveries that would have dramatically found shortly afterward in the Southwest was the closely
altered our view and understanding of dinosaurs, birds, related but much larger crested theropod Dilophosaurus of
and their evolution, but that event would have to wait an- the Early Jurassic.
other three-quarters of a century. What really spurred the science of dinosaur research
The mistake of the American Museum expeditions in were the Yale expeditions to Montana in the early 1960s
heading northeast contributed to a set of problems that that dug into the little-investigated Early Cretaceous Clo-
seriously damaged dinosaur paleontology as a science verly Formation. The discovery of the Velociraptor relative
between the twentieth-century world wars. Dinosaurol- Deinonychus finally made it clear that some dinosaurs were
ogy became rather ossified, with the extinct beasts widely sophisticated, energetic, agile dinobirds, a point reinforced
portrayed as sluggish, dim-witted evolutionary dead ends by the realization that they and the other sickle claws, the
doomed to extinction, an example of the “racial senes- troodontids, as well as the ostrichlike ornithomimids, had
cence” theory that was widely held among researchers who fairly large, complex brains. These developments led John
preferred a progressive concept of evolution at odds with Ostrom to note and detail the similarities between his Dei-
more random Darwinian natural selection. It did not help nonychus and Archaeopteryx and to conclude that birds are
matters when artist-paleontologist Gerhard Heilmann the descendants of energetic small theropod dinosaurs. Os-
published a seminal work that concluded that birds were trom further observed that the bird- and mammal-like body
not close relatives of dinosaurs, in part because he thought design of dinosaurs meant that that their newly apparent
dinosaurs lacked a wishbone furcula, which had just been presence in Arctic regions could not be taken as evidence
found, but misidentified, in Oviraptor. The advent of the that those ancient polar habitats were warm even in winter.
Depression, followed by the trauma of World War II— Realizing that the consensus dating back to the origi-
which led to the loss of some important fossils on the con- nal discovery that dinosaurs were an expression of the
tinent as a result of Allied and Axis bombing—brought reptilian pattern was flawed, Robert Bakker in the 1960s
major dinosaur research to a near halt. and 1970s issued a series of papers contending that dino-
Even so, public interest in dinosaurs remained high. saurs and their feathered descendants constituted a dis-
The paleoart of Charles Knight made him famous. The tinct group of archosaurs whose biology and energetics
Star Wars–Jurassic Park of its time, RKO’s King Kong of 1933 were more avian than reptilian. In the eye-popping arti-
amazed audiences with its dinosaurs seemingly brought to cle “Dinosaur Renaissance” in a 1975 Scientific American,
life. Two major film comedies, 1938’s Bringing Up Baby, Bakker proposed that some small dinosaurs themselves
starring Cary Grant and Katherine Hepburn, and 1949’s were feathered. In the late 1970s, Montana native John
On the Town, featuring Gene Kelly and Frank Sinatra, in- Horner found baby hadrosaurs and their nests, providing
volved climactic scenes in which sauropod skeletons at a the first look at how some dinosaurs reproduced. At the
semifictional New York museum collapsed because of the same time, researchers from outside paleontology stepped
hijinks of the lead characters. Unfortunately, the very pop- into the field and built up the evidence that the impact of
ularity of dinosaurs gave them a circus air that convinced a mountain-sized asteroid was the long-sought great dino-
many scientists that they were beneath their scientific dig- saur killer. This extremely controversial and contentious
nity and attention. idea turned into the modern paradigm when a state-sized
Despite the problems, discoveries continued. In an meteorite crater was found in southeastern Mexico dating
achievement remarkable for a nation ravaged by the Great to the end of the dinosaur era. At the same time, the vol-
Patriotic War and suffering under the oppression of Stalin- canic Deccan Traps of India were proposed as an adjunct
ism, the Soviets mounted postwar expeditions to Mongolia in the demise of the nonbird dinosaurs.
that uncovered the Asian version of Tyrannosaurus, a.k.a. These radical and controversial concepts greatly boosted
Tarbosaurus, and the enigmatic arms of enormous clawed popular attention on dinosaurs, culminating in the Jurassic
Therizinosaurus. Equally outstanding was how the Poles, Park novels and films that sent dinomania to unprecedented
led by the renowned Halszka Osmólska, took the place heights. The elevated public awareness was combined with
of the Soviets in the 1960s, discovering in the process the digital technology in the form of touring exhibits of ro-
famed complete skeleton of Velociraptor engaged in combat botic dinosaurs. This time the interest of paleontologists
with Protoceratops. They too found another set of mysteri- was raised as well, inspiring the second and ongoing golden
ous long arms with oversized claws, Deinocheirus. Another age of dinosaur discovery and research, which is surpass-
leader of Mongolian paleozoology has been the native Rin- ing that which has gone before. Assisting the work are im-
chen Barsbold. proved scientific techniques in the area of evolution and
In the United States, Roland Bird studied the track- phylogenetics, including cladistic genealogical analysis,
ways of herds of Texas-sized Cretaceous sauropods before which has improved the investigation of dinosaur relation-
World War II. Shortly after the global conflict, the Triassic ships. New generations of artists have portrayed dinosaurs
Ghost Ranch quarry in the Southwest, packed with com- with a “new look” that lifts tails in the air and gets feet off
plete skeletons of little Coelophysis, provided the first solid the ground to represent the more dynamic gaits that are in
10
History of Discovery and Research
Deinonychus and
Tenontosaurus
line with the more active lifestyles the researchers now rising economies of many second-world nations, reducing
favor. This artist and researcher noticed that the sickle- the need to import Western expertise.
clawed dromaeosaurs and troodonts, as well as the ovirap- In South America, Argentine and American paleontolo-
torosaurs, possessed anatomical features otherwise found gists collaborated in the 1960s and 1970s to reveal the first
in flightless birds and suggested that these dinosaurs were Middle and Late Triassic protodinosaurs, finally showing
also secondarily flightless. that the very beginnings of dinosaurs started among sur-
Dinosaurs are being found and named at an unprec- prisingly small archosaurs. Since then, Argentina has been
edented rate as dinosaur science goes global, with efforts the source of endless remains from the Triassic to the end
under way on all continents. In the 1970s the annual So- of the Cretaceous that include the early dinosaurs Eoraptor
ciety of Vertebrate Paleontology meeting might have seen and Herrerasaurus, supertitanosaur sauropods such as Argen-
a half-dozen presentations on dinosaurs; now it is in the tinosaurus, Futalognkosaurus, and Dreadnoughtus, and oversized
area of a couple of hundred. Especially important has been theropods such as Giganotosaurus that preyed on them.
the development of local expertise made possible by the Among the most extraordinary finds have been sauropod
11
History of Discovery and Research
nesting grounds that allow us to see how the greatest land preserved material as well as of strife, as the locals con-
animals of Earth’s history reproduced themselves. tend with the authorities for the privilege of excavating
In southern Africa excellent remains of an Early Juras- the fossils for profit—sometimes altering the remains to
sic species of Coelophysis verified how uniform the dinosaur “improve” them—rather than for rigorous science. The
fauna was when all continents were gathered into Pangaea. feathered dinosaurs soon included the potential ovirap-
Northern Africa has been the major center of activity as a torosaur Caudipteryx. Even more astonishing have been
host of sauropods and theropods have filled in major gaps the Yixian-Jiufotang dromaeosaurs. These small sickle
in dinosaur history. Australia is geologically the most sta- claws bear fully developed wings not only on their arms
ble of continents, with relatively little in the way of tectoni- but on their similarly long legs as well. This indicates not
cally driven erosion to either bury fossils or later expose only that dromaeosaurs first evolved as fliers but that they
them, so dinosaur finds have been comparatively scarce were adapted to fly in a manner quite different from the
despite the aridity of the continent. The most important avian norm. The therizinosaur Beipiaosaurus looks like a
discoveries have been of Cretaceous dinosaurs that lived refugee from a Warner Brothers cartoon. But the Yixian-
close to the South Pole, showing the climatic extremes di- Jiufotang is not just about confirming that birds are dino-
nosaurs were able to adapt to. Glacier-covered Antarctica saurs and that some dinosaurs were feathered. One of the
is even less suitable prospecting territory, but even it has most common dinosaurs of the Early Cretaceous was the
produced the Early Jurassic crested avepod Cryolophosaurus parrot-beaked Psittacosaurus. Although it was known from
as well as other dinosaur bones. numerous skeletons across Asia found over the last eighty
At the opposite end of the planet, the uncovering of a years, no one had a clue that its tail sported large, arcing,
rich Late Cretaceous fauna on the Alaskan North Slope bristly spines until a complete individual with preserved
confirms the ability of dinosaurs to dwell in latitudes cold skin was found in the Yixian-Jiufotang. To top things off,
and dark enough in the winter that lizards and crocodilians the Yixian-Jiufotang has produced the small ornithischian
are not found in the same deposits. Farther south, a cadre Tianyulong, which suggests that insulating fibers were wide-
of researchers have continued to plumb the great dinosaur spread among small dinosaurs. There are new museums in
deposits of western North America as they build the most China brimming with enormous numbers of undescribed
detailed sample of dinosaur evolution from the Triassic dinosaur skeletons on display and in storage. Making the
until their final loss. We now know that armored ankylo- Yixian-Jiufotang deposits all the more interesting is that
saurs were roaming along with plated stegosaurs in the they appear to be the rare remnants of highland lakes
Morrison Formation, a collection of sauropods has been thousands of meters above sea level, where chilling win-
exposed from the Early Cretaceous, and one new ceratop- ters were similar to those at the poles. Lately coming from
sian and hadrosaur and theropod and ankylosaur after an- central Asia are the cartoonish dromaeosaurs Halszkarap-
other is coming to light in the classic Late Cretaceous beds. tor and Natovenator, which may have swum for their prey
Now Mongolia and especially China have become a great and grabbed at it with their duck-like heads.
frontier in dinosaur paleontology. Even during the chaos On a global scale, the number of dinosaur trackways
of the Cultural Revolution, Chinese paleontologists made that have been discovered is in the many millions. This
major discoveries, including the first spectacularly long- is logical in that a given dinosaur could potentially con-
necked mamenchisaur sauropods. As China modernized tribute only one skeleton to the fossil record but could
and Mongolia gained independence, Canadian and Amer- make innumerable footprints. In a number of locations,
ican researchers worked with their increasingly skilled and trackways are so abundant that they form what have been
independent resident scientists, who have become a lead- called “dinosaur freeways.” Many of the trackways were
ing force in dinosaur research. It was finally realized that formed in a manner that suggests their makers were mov-
the oviraptors found associated with nests at the Flaming ing in herds, flocks, packs, and pods. A few may record the
Cliffs were not eating the eggs but brooding them in a prea- attacks of predatory theropods on herbivorous dinosaurs.
vian manner. Almost all of China is productive when it The history of dinosaur research is not just one of new
comes to dinosaurs, and after many decades paleontolo- ideas and new locations; it is also one of new techniques
gists started paying attention to the extraordinary fossils and technologies. The turn of the twenty-first century has
being dug up by local farmers from Early Cretaceous Yix- seen paleontology go high tech with the use of computers
ian-Jiufotang lake beds in the northeast of the nation. for processing data and high-resolution CT scanners to peer
In the mid-1990s, complete fossils of small compsog- inside fossils without damaging them. Skeletal and life res-
nathid theropods labeled Sinosauropteryx began to show torations are being generated in 3-D digital format—
up with their bodies covered with dense coats of bristly whether this has improved the process is another matter.
protofeathers, to the delight of Ostrom and others. More Dinosaurology has also gone microscopic and molecular in
recently it has been argued that it is often possible to deter- order to assess the lives of dinosaurs at a more intimate
mine the color of the feathers! This was just the start: the level, telling us how fast they grew, how long they lived, and
Yixian-Jiufotang beds are so extensive and productive that at what age they started to reproduce. Bone isotopes are
they have become an inexhaustible source of beautifully being used to help determine dinosaur diets and to state that
12
History of Discovery and Research
some dinosaurs were semiaquatic. And it turns out that unlikely that a reorganization of similar scale will occur in
feather pigments can be preserved well enough to restore the future, but we now know enough about the inhabitants
original colors. Meanwhile the Jurassic World franchise helps of the Mesozoic to have the basics well established. Sauro-
sustain popular interest in the group even as it presents an pods will not return to a hippo-like lifestyle, and dinosaurs’
obsolete, prefeather image of the birds’ closest relations. tails will not be chronically plowing through Mesozoic
The evolution of human understanding of dinosaurs has muds. Dinosaurs are no longer so mysterious. Even so, the
undergone a series of dramatic transformations since they research is nowhere near its end. To date, over 800 valid
were scientifically discovered almost two hundred years dinosaur species in about 500 genera have been discovered
ago. This is true because dinosaurs are a group of “exotic” and named. This probably represents at most a quarter,
animals whose biology was not obvious from the start, un- and perhaps a much smaller fraction, of the species that
like fossil mammals or lizards. It has taken time to build up have been preserved in sediments that can be accessed.
the knowledge base needed to resolve their true form and And, as astonishingly strange as many of the dinosaurs
nature. The latest revolution is still young. When I was a uncovered so far have been, there are equally odd species
youth, I learned that dinosaurs were, in general, sluggish, waiting to be unearthed. Reams of work based on as-yet-
cold-blooded, tail-dragging, slow-growing, dim-witted rep- undeveloped technologies and techniques will be required
tiles that did not care for their young. The idea that some to provide further details about both dinosaur biology and
were feathered and that birds are living descendants was the world in which they lived. And although a radical new
beyond imagining. Dinosaurology has matured in that it is view is improbable, there will be many surprises.
13
What Is a Dinosaur?
WHAT IS A DINOSAUR?
To understand what a dinosaur is, we must first start higher femur is a cylinder turned in at a right angle to the shaft of
in the scheme of animal classification. The Tetrapoda are the femur that fits into a cylindrical, internally open hip
the vertebrates adapted for life on land—amphibians, rep- socket. This allows the legs to operate in the nearly verti-
tiles, mammals, birds, and the like. Amniota comprises cal plane characteristic of the group, with the feet directly
those tetrapod groups that reproduce by laying hard-shelled beneath the body. You can see this system the next time
eggs, with the proviso that some have switched to live birth. you have chicken thighs. The ankle is a simple fore-and-
Among amniotes are two great groups. One is the Synap- aft hinge joint that also favors a vertical leg posture. Dino-
sida, which includes the archaic pelycosaurs, the more ad- saurs were “hind-limb dominant” in that they were either
vanced therapsids, and mammals, which are the only bipedal or, even when they were quadrupedal, most of the
surviving synapsids. The other is the Sauropsida, much the animal’s weight was borne on the legs, which were always
same as the Reptilia, all but some early forms of which be- built more strongly than the arms. The hands and feet
long to the great Diapsida, which included most of the an- were generally digitigrade, with the wrist and ankle held
cient marine reptiles that are detailed in the Princeton field clear of the ground. All dinosaurs shared a trait also wide-
guide on that set of creatures. Surviving diapsids include spread among archosaurs in general, the presence of large
turtles, the lizard-like tuataras, true lizards and snakes, and often remarkably complex sinuses and nasal passages.
crocodilians, and birds. The Archosauria is the largest and Aside from the above basic features, dinosaurs, even
most successful group of diapsids and includes crocodilians when we exclude birds, were an extremely diverse group
and dinosaurs. Birds are literally flying dinosaurs. of animals, rivaling mammals in this regard. Dinosaurs
Archosaurs also include the basal forms informally ranged in form from nearly birdlike types such as the
known as thecodonts because of their socketed teeth, them- sickle-clawed dromaeosaurs to rhino-like horned ceratop-
selves a diverse group of terrestrial and aquatic forms that sians to armor-plated stegosaurs to elephant- and giraffe-
include the ancestors of crocodilians and the winged ptero- like sauropods and dome-headed pachycephalosaurs.
saurs (covered in another Princeton field guide), although They even took to the skies in the form of birds. However,
those fliers were not intimate relatives of dinosaurs and dinosaurs were limited in that they were persistently terres-
birds. trial. Although some dinosaurs may have spent some time
The great majority of researchers now agree that the feeding in the water like moose or fishing cats, at most a
dinosaurs were monophyletic in that they shared a com- few became strongly amphibious in the manner of hippos,
mon ancestor that made them distinct from all other ar- much less marine like seals and whales. The only strongly
chosaurs, much as all mammals share a single common aquatic dinosaurs are some birds. The occasional state-
ancestor that renders them distinct from all other synap- ment that there were marine dinosaurs is therefore very
sids. This consensus is fairly recent—before the 1970s it incorrect—these creatures of Mesozoic seas were various
was widely thought that dinosaurs came in two distinct forms of reptiles that had evolved over the eons.
types that had evolved separately from thecodont stock, The situation within dinosaurs is interesting because it
the Saurischia and Ornithischia. It was also thought that has become chaotic. For over a century, dinosaurs have
birds had evolved as yet another group independently from generally been segregated into the two classic groups: the
thecodonts. Dinosauria is formally defined as the phyloge- Saurischia, which includes the largely herbivorous sau-
netic clade that includes the common ancestor of Tricer- ropodomorphs and the usually predaceous theropods, birds
atops and birds and all their descendants. Because different included; and the vegetarian Ornithischia, which despite the
attempts to determine the exact relationships of the ear- name has nothing to do with birds. But while ornithischi-
liest dinosaurs produce somewhat different results, there ans are clearly a real, monophyletic clade, the saurischian
is some disagreement about whether the most archaic, group has always been weak. Part of the problem is that
four-toed theropods were dinosaurs or lay just outside the the earliest species of a given group are all pretty similar to
group. This book includes them, as do most researchers. one another, and it is difficult to parse out the few, still sub-
In anatomical terms, one of the features that most distin- tly different characters that show they are at the base of one
guish dinosaurs centers on the hip socket. The head of the of the greater, more sophisticated groups but not the other.
A basal archosaur,
Euparkeria
14
Dating Dinosaurs
DATING DINOSAURS
How can we know that dinosaurs lived in the Mesozoic, ago. Deposited largely by rivers over an area covering
first appearing in the Late Triassic over 230 million years many states in the continental interior, the Morrison For-
ago and then disappearing at the end of the Cretaceous 66 mation is easily distinguished from the marine Sundance
million years ago? Formation lying immediately below as well as from the
As gravels, sands, and silts are deposited by water and similarly terrestrial Cedar Mountain Formation above,
sometimes wind, they build up in sequence atop the previ- which contains very different sets of dinosaurs. Formed
ous layer, so the higher in a column of deposits a dinosaur over millions of years, the Morrison is subdivided into
is, the younger it is relative to dinosaurs lower in the sed- lower (older), middle, and upper (younger) levels. So a fos-
iments. Over time, sediments form distinct stratigraphic sil found in the Sundance is older than one found in the
beds that are called formations. For example, Apatosaurus, Morrison, a dinosaur found in the lower Morrison is older
Brontosaurus, Diplodocus, Barosaurus, Stegosaurus, Camptosau- than one found in the middle, and a dinosaur from the
rus, Allosaurus, and Ornitholestes are found in the Morri- Cedar Mountain is younger still.
son Formation of western North America, which was laid Geological time is divided into a hierarchical set of
down in the Late Jurassic, from 150 to 156 million years names. The Mesozoic is an era—preceded by the Paleozoic
15
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