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 Praise for
The Lost World of the Dinosaurs

“An insightful and informative meander through the evolution of dinosaurs

and other extinct species, with a touch of personal flair. Armin Schmitt has
a passion for dinosaurs, which you can feel in his writing.”
—Steve Brusatte, professor and paleontologist at the University of Edinburgh and New York
Times bestselling author of The Rise and Fall of the Dinosaurs

“Through personal stories from fieldwork in the far corners of the world to
the latest discoveries in research laboratories, Armin Schmitt traces the
history of the dinosaurs, their evolution, and their environment. In The Lost
World of the Dinosaurs, you’ll encounter issues as diverse as the bizarre
marine reptiles of the deserts of Nevada, the ethical questions of the
scientific study of Myanmar amber, and why Tyrannosaurus rex deserves
its reputation as the most beloved dinosaur. Schmitt’s book will delight
those new to paleontology, and even dinosaur devotees will learn something
new.”
—Thomas R. Holtz Jr., principal lecturer in vertebrate paleontology at the University of
Maryland

“An enthusiastic survey of the ancient world. Schmitt’s tale blends memoir,
science history, and new discoveries into an exuberant celebration of
Mesozoic life.”
—Riley Black, author of When the Earth Was Green and The Last Days of the Dinosaurs

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ARMIN SCHMITT has been fascinated by dinosaurs since he was five
years old. During his studies in paleontology at Bonn, he also rose to the
position of collection manager at the university’s Goldfuß-Museum. Most
recently, he was a research assistant at the Department of Earth Sciences at
the University of Oxford. Today, he is a vertebrate paleontologist as well as
a PhD student at the University of Cambridge.

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The Lost World of the Dinosaurs
Uncovering the Secrets of the Prehistoric Age

Armin Schmitt
With illustrations by Ben Rennen

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For Elisabeth and Maximilian

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 Contents

Prologue
The Great Dying

The Triassic (251.9 to 201.3 million years before present)

Image

Chapter 1: New Life in the Sea

Omphalosaurus: Crushing It!

Nevada’s Ichthyosaurs: All Just One Big Family
Of Strange Monsters and Underwater Flight

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Chapter 2: New Life on Land

Survival in Hell: The Road to Dinosaur Domination

The Dragon Who Wanted to Be a Tyrannosaurus

Where the Wild Things Are

The First Giants: Hundreds of Long-Necked Dinosaurs Stuck

in Mud
The Jurassic (201.3 to 145 million years before present)

Image
Chapter 3: The Dinosaurs of the Morrison Formation

The Bone Wars

Chapter 4: Germany’s Dinosaurs
 How, on an Island, a Giant Turned into a Dwarf
The First Feather That Changed Everything: Archaeopteryx

The Adventures of Werner Janensch

The Lower Cretaceous (145 to 100.5 million years before present)

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Chapter 5: Argentina—Where the Giants Live

The Long-Necked Dinosaurs That Tower over Everything

Chapter 6: Myanmar—Trapped in “Liquid Gold”

Feathers, Claws, and Ticks in Amber
A Dinosaur Even Smaller than a Hummingbird?

A Moral Dilemma

Chapter 7: The Spinosaurs

The Adventures of Ernst Stromer in Egypt

Spinosaurus: On Land, on the Shore, or in the Water?

Fish and No Chips: The Fish Eaters of the Isle of Wight

Chapter 8: More than Just Bones

The Super-Lung

The Cloaca: How Do Dinosaurs Make Wee-Wee?

Do Female Dinosaurs Have Ovaries?

The Grace of the Sleeping Club-Tail Dragon

The Upper Cretaceous (100.5 to 66 million years before present)

Image
Chapter 9: Wyoming—The Hell Creek

Triceratops: A Tank on Four Legs

How a Cretaceous Landscape is Resurrected
Chapter 10: Tyrannosaurus—The Measure of All Things!

The Power of the Brand: Why Tyrannosaurus Is So Popular

A Record Sum for the King

How Many Tyrannosaurs Were There?

Why Tyrannosaurus Was Not a Scavenger

Different Prey for Father and Son

I Am the King and Not a Peacock: Did T. rex Have Feathers?

Short Arms Are Normal but Long Ones Are Strange

Boy or Girl: Was Sue Female?

The King, the Queen, and the Emperor

Chapter 11: Movement Captured in the Stone: What Footprints Tell
Us

The German Dinosaur Highway

The Dinosaur Tracks of Lommiswil

The Track Site of Cal Orck’o in Bolivia

Chapter 12: Birds—The Last Dinosaurs

Where Is the First Ostrich?

Almost Like Easter: An Egg Thief Who Lays Colorful Eggs

The Goddess Asteria: The Mother of All Chickens and Geese
 Beyond the Rainbow

Chapter 13: The End of the Dinosaurs

The Day it Rained Glass
Like a Phoenix From the Ashes: How Birds Survived the
Catastrophe

Epilogue

Glossary

References

Acknowledgments
Phylogeny Chart

Time Scale

Index

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 Prologue

Since the age of five, I have been fascinated by the giant dinosaurs of
prehistoric times. I am very grateful that my passion and purpose were
revealed to me so early on and that I was able to turn this enthusiasm for
dinosaurs into my profession. Today, when I find a dinosaur bone in the
field, I still have the same glow in my eyes as I did as a small child when I
first saw real skeletons of these giants at the Senckenberg Natural History
Museum in Frankfurt. The path from dino-crazy child to vertebrate
paleontologist, however, was not straightforward by any stretch of the
imagination. Nevertheless, I had the great fortune to be part of many
discoveries, some of them very important, even if I myself was not at the
center of the research. Some research projects I was involved in contributed
to a better understanding of the evolution of dinosaurs and other prehistoric
reptiles and fundamentally changed our view of dinosaurs. Some of that is
reflected in this book. It’s a great feeling to be at the heart of research in the
golden age of paleontology, using cutting-edge technology to uncover new
mysteries and unravel puzzles that just a few years ago were thought never
to be explained. One reason we’ve recently learned so many new things
about dinosaurs is the incredible rate at which new dinosaur species are
being scientifically described. In the last twenty years alone, more dinosaur
species have been discovered per year than ever before. Since 2003, about
forty-five new dinosaurs have been discovered each year. This is mainly
due to increased funding for research, more scientists today, and more
teams in the field. Remote sites are more accessible, and we have
increasingly accurate geological maps with precise coordinates for these
sites. It is not uncommon for new fossils to be discovered at already known
localities, and it is not uncommon for this to bring new species to light.
Also partly responsible is the rapid increase in computing power and speed
of computers and workstations, which make it possible to generate
simulations and three-dimensional models and to evaluate complex
statistics to generate phylogenetic trees of dinosaurs. And recently
discovered dinosaur species not only fill museums and provide new names
that are difficult to pronounce, but also provide insights into how different
animals were related as well as their habitats and survival strategies. Each
new find gives us ever-more-precise insights into the ecosystem of the
dinosaurs, and their external appearance even helps us better understand
their internal structure and the function of their organs. This is enormously
important because some dinosaurs had such unusual anatomies that we
cannot find anything comparable in nature today. New discoveries help us
understand how dinosaurs were able to cope with drastic climate changes
and adapt to new habitats. Dinosaurs were fascinating creatures that can
serve as model organisms to understand many mechanisms of nature and
evolution. In this book, based on several prominent examples of species of
dinosaurs, I discuss the fact that there are always new discoveries to be
made and that, despite our extensive knowledge of dinosaurs, many
questions remain unanswered.

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 The Great Dying

When the Earth was burning and the sea was choking

To understand how dinosaurs became the most successful terrestrial

vertebrate group in Earth’s history, we must travel to the beginnings of
dinosaur evolution and clarify what factors were necessary for dinosaurs to
begin their triumphant reign.
Most children today know that all dinosaurs, with the exception of
modern birds, became extinct at the end of the Cretaceous period about 66
million years ago. An asteroid hit Earth at that time and sealed the fate of
the dinosaurs and many other animal groups. Ammonites disappeared from
the oceans.

These were cephalopods related to today’s Nautilus and squids. They

lived in our planet’s oceans for well over 300 million years and were an
important component of complex food webs in marine ecosystems. Along
with the ammonites, large marine reptiles such as the four-flippered
plesiosaurs and the varanid-like mosasaurs also disappeared without any
descendants.
 But as terrible as this cosmic impact must have been, it is dwarfed by an
event that had much greater consequences and is widely considered the
worst catastrophe for life on this planet. It was a cataclysm of apocalyptic
proportions, so devastating that some authors refer to it as “the Great
Dying”: the Permian–Triassic mass extinction.
Michael Benton, one of the most influential paleontologists of our time,
wrote a book about it, giving it the striking title When Life Nearly Died.
This event, which took place at the end of the Permian period, resulted in
the greatest mass extinction event in the history of the Earth. About 90
percent of all species living at that time became extinct. For the first time in
Earth’s history, many insect species were also affected.
During the Permian, food was still plentiful for all animals. The Earth
was a green planet. For millions of years, plant growth had been so
abundant throughout the world that the oxygen content in the atmosphere
continued to rise and reached its highest level in the entire history of the
Earth. Today, we assume that the atmospheric oxygen level was around 30
percent in the Permian.

By comparison, the oxygen level in the atmosphere today is

only about 21 percent.

Droughts, fires, air pollution, and acid rain subsequently destroyed the
lush forests, causing the atmospheric oxygen content to drop from 30 to
about 10 to 15 percent and then remain at low levels for many millions of
years. Oxygen levels would never again reach those from the beginning of
the Permian. The effects were so far-reaching that it took almost 15 million
years for the Earth’s forests to fully recover.
“The Great Dying,” and the subsequent collapse of the ecosystems, is
thought to have been caused by mega-volcanism in Russia. The so-called
Siberian traps, or flood basalts, are a testimony to this catastrophe. These
are extensive lava flows that stretch from the Barents Sea in the northwest
to Kazakhstan in the southwest. From there, they continue to Novosibirsk
and Irkutsk in the south. In the east, they extend beyond the Lena River
almost to Yakutsk. Some of you may recognize Irkutsk and Yakutsk from
the board game Risk. The volcanic eruptions occurred in three spurts, over a
period of about nine hundred thousand years, and covered an area of 7
million square kilometers with volcanic rock. This is equivalent to almost
the entire area of Australia!
The volcanic eruptions responsible for this are considered to be among
the largest known volcanic events in Earth’s history. The eruptions took
place about 252 million years ago at the Permian–Triassic boundary and
their consequences are causally linked to the mass extinction at the end of
the Permian, or “the Great Dying.” The postapocalyptic world after the
Permian catastrophe must have come very close to our idea of hell. The
mega-volcanism in Siberia released substantial amounts of carbon dioxide,
fluorine, hydrogen chloride, and sulfur dioxide, which reacted with rain to
form sulfuric acid, permanently damaging or destroying both marine and
terrestrial habitats. Carbon dioxide emissions triggered global warming,
increasing the global temperature by 9°F in a very short period of time. We
now also suspect that the ozone layer was severely damaged by the
greenhouse gases and the fly ash, resulting in an increase in UV radiation,
in turn hampering plant growth and causing pollen malformations. Massive
forest fires accelerated the large-scale vegetation loss, and acid rain finished
off most of the plants. At that time, it only rained periodically, during the
mega-monsoon season. But then, the celestial floodgates would open wide
and pour forth torrents of caustic rain for days and weeks across the
supercontinent of Pangaea. The rest of the year was marked by terrible heat
waves, droughts, fires, and an unbearable average temperature of 113°F.
While there was still sparse vegetation in the coastal regions, the interior
landmass was dominated by vast deserts wider than even the Sahara or
Gobi deserts. The consequences were devastating. The extreme
environmental conditions increased erosion on land. This resulted in
extensive draining of the soil into the sea, leading to eutrophication effects
such as algal blooms in marine habitats. At the same time, there was a mass
proliferation of marine protozoa in oxygen-free environments, whose
metabolic products—in addition to all the gases from the massive
volcanism—released methane, halocarbons, and large amounts of hydrogen
sulfide (H2S) into the atmosphere.
In the meantime, anoxic—oxygen-free—conditions developed in the
oceans, causing all reefs and their inhabitants to die off. Rocks that bear
witness to this momentous event are called black shales, which are dark-
colored organic-rich sediments. With the formation of anoxic marine zones,
the rapid decrease in pH, and the release of methane hydrate, a mass
extinction began in the oceans. As a result of the methane escaping into the
atmosphere, the Earth’s temperature rose by a further 9°F and the upper
water layers of the world’s oceans even warmed by at least 14°F. The Earth
was burning and the oceans were boiling. The ever-increasing concentration
of greenhouse gases led to a domino effect, which we describe as a
galloping greenhouse effect, that explains why the mega-volcanism in
Siberia lasted almost a million years but the mass extinction only thirty
thousand. Initially, the changes were gradual, but as more and more habitats
were damaged and more and more pollutants were emitted, temperatures
increased faster and faster, and the collapse of the ecosystems became
unstoppable. In the end, about 95 percent of all marine species and about 75
percent of all terrestrial species became extinct during this event. Among
the few survivors were the direct ancestors of dinosaurs and crocodiles: the
so-called archosaurs.
Without this catastrophe, the triumphant rise of the dinosaurs would
probably never have happened. And so it is perhaps an irony of fate that, on
the one hand, the dinosaurs entered the world stage after a devastating mass
extinction, while on the other, they then fell victim to a mass extinction
themselves. There have been five such events of mass extinction in the
history of the Earth. Three of them are closely related to the origin of the
dinosaurs and were crucial for their unprecedented evolutionary success—
but ultimately also responsible for their demise:

1. The Permian catastrophe about 252 million years ago (“the

Great Dying”) favored the development of the archosaurs. The
archosaurs, or “ruling lizards,” include all crocodiles and
dinosaurs. Birds also belong to this group, but they did not yet
exist in the Triassic. The archosaurs, and dinosaurs in
particular, were able to hold their own well in the hostile world
of the Triassic because their efficient lungs were better able to
cope with the low levels of atmospheric oxygen, they could run
faster with their long, straight legs, and their bodies were
farther from the hot ground than those of other lizards and
amphibians. These traits were advantageous in the oftentimes
desertlike regions of the prehistoric continent. This allowed
them to colonize new habitats. Their scaly skin prevented their
 bodies from drying out, which was an invaluable advantage in
the sweltering heat of the Triassic.
2. The end-Triassic mass extinction, which occurred about 201
million years ago. This catastrophe killed the conodonts in the
sea, which had existed for about 340 million years and now
disappeared without descendants. As either the first vertebrates,
or close relatives of today’s vertebrates, they are among the
earliest forms that later gave rise to fish, amphibians, reptiles,
mammals, and birds. Further impacting dinosaur evolution was
the extinction of four major groups of crocodilian relatives at
the end of the Triassic: rauisuchians, phytosaurs, aetosaurs, and
ornithosuchians. These representatives of the crocodilian
lineage were similarly as successful as the dinosaurs in the
Triassic. At the same time, almost all temnospondyls, a clade of
giant amphibians that could grow up to thirteen feet long,
became extinct. They are considered a sister group of today’s
frogs and salamanders. For simplicity, they can be thought of
as thirteen-foot-long monster salamanders with bony plates on
their skin. With the demise of the monster salamanders and
most land-dwelling crocodiles, there was no longer any
competition for the dinosaurs, and so they became the
undisputed rulers of the Earth at the beginning of the Jurassic.
3. The end-Cretaceous mass extinction about 66 million years ago
finally sealed the fate of all dinosaurs that were not true birds
and not on the bird lineage. The end of this book explores why
the dinosaurs did not survive the asteroid impact.
 The fossil record includes all scientifically documented finds
of fossils worldwide. The occurrence of fossils at a certain
place is usually mentioned in scientific publications. Therein,
the geographical location as well as the stratigraphic context
of the finds are given. Investigating, adding to, and ultimately
completing the fossil record are some of the most important
tasks of paleontology.

Between these three catastrophes stretched incredibly long periods of

time. The evolution of the dinosaurs took place between the Permian
catastrophe and the impact of the asteroid at the end of the Cretaceous
period. These events are about 186 million years apart. In order to imagine
such a time span, one must keep in mind that Tyrannosaurus and humans
are separated by a little more than 65 million years. The last Tyrannosaurus
became extinct 66 million years ago and humans (Homo sapiens) emerged
about three hundred thousand years ago. Stegosaurus, on the other hand,
became extinct almost 148 million years ago, so it lived about 80 million
years before Tyrannosaurus. So, in terms of geologic time, these two are
further apart than Tyrannosaurus and humans. But the very first dinosaurs
emerged another 90 million years or so before Stegosaurus. Consequently,
Plateosaurus (an ancestor of the later long-necked dinosaurs) lived in a
completely different world than Stegosaurus, and Tyrannosaurus, in turn,
lived in a completely different world than Stegosaurus. The reign of the
dinosaurs lasted so incredibly long that entire continents broke up, seas
opened and closed again, and entire oceans were created during it.
This thrilling journey of the dinosaurs was sandwiched between two of
Earth’s greatest mass extinction events. In both cases, these were global
disasters. Why is this important? Dinosaurs were the most successful land
creatures in the long history of the Earth. No other group of animals reigned
on land as long as they did. Yet, at the end of the Cretaceous period, they
fell victim to a mass extinction triggered by changes in climate and
vegetation. While the cause of this extinction was the asteroid, it only killed
a few dinosaurs directly. Countless more animals died in agony in the
weeks and months following the impact, after the sky darkened,
temperatures changed, forests died, and the food chain collapsed.
This scenario sounds familiar. Today, man is certainly the most
successful complex organism on Earth. Just like the dinosaurs, we have
colonized every continent, hold our own against every enemy, and use all of
the Earth’s resources—at the same time, the fastest climate change we
know of in all of Earth’s history is taking place around us, while we face
unprecedented species extinctions, the scope of which cannot yet be
foreseen. According to a 2019 estimate, the number of birds in North
America has declined by about 3 billion since 1970, or about 30 percent of
the population there. In Germany, according to the Nature and Biodiversity
Conservation Union, or NABU, the number of breeding pairs of birds
declined from 97.5 million in 1998 to 84.8 million in 2009, a decrease of 15
percent in just eleven years. Never before in Earth’s history have so many
birds died. At the same time, the number of flying insects in Germany’s
nature reserves has declined by an extremely alarming 76 percent over the
past twenty-seven years. But while we might be glad that there are fewer
mosquitoes at night, flying insects are the main pollinators of most crops
and at the same time an important contributor to the diet of most bird
species.
The causes of climate change at the end of the Cretaceous period are
different than today, but that does not make the current situation any less
severe. The extinction of the dinosaurs had extrinsic causes, while
anthropogenic influences are responsible for our current situation. Humans
themselves are driving climate change and species extinction.
I often hear that some people don’t believe in climate change, and don’t
believe that humans have any significant impact on climate change. The
devastating species loss we are currently experiencing makes surprisingly
few headlines. Yet climate change and species extinction can be observed in
nature. They can be objectively measured and proven with reproducible
scientific experiments, whether you believe in them or not. Personal views
and opinions are irrelevant here. And if we choose to ignore the warning
signs, the problems will not go away—they will only get worse.

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 The Triassic

(251.9 to 201.3 million years

before present)

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OceanofPDF.com
 Chapter 1

New Life in the Sea

Omphalosaurus: Crushing It!

It is morning on Monday, September 26, 2011, and I am climbing up to our
dig site from base camp at the foot of the Augusta Mountains in Nevada, as
I have been doing for several days. It is about sixteen hundred feet above
the camp and there are no roads or footpaths leading to it, only a trail left by
wild mustangs. Today the ascent is particularly arduous, because I’m
carrying a bag of plaster up the mountain in addition to provisions, photo
equipment, and 1.25 gallons of water. The first few miles we could still
climb in the shade of the mountain, but by the time we reach the first
plateau, the hot sun burns down on us. There is no more shade, because
there are no trees growing up here, anywhere. Since it is too dry in the
valley, only sagebrush and grasses grow, and their barbs stick to stockings,
shoes, and clothing and irritate the skin. In the Augusta Mountains, juniper
bushes do not appear until about five thousand feet above sea level, and
well above sixty-five hundred feet, almost at the summit of Cain Mountain,
grow scattered bony, gnarled pinyon pines and the so-called Mormon tea
(ephedra). This phenomenon is called the inverse tree line. In the Alps it is
the other way around—there, the trees grow in the valley and the higher
you go up, the rarer they become, until—between fifty-nine hundred and
seventy-two hundred feet above sea level—they disappear altogether.
From the first plateau you can still see our camp and communicate with it
via walkie-talkie. In 2011, there was no cell phone reception here. But you
had no electricity to charge your cell phone anyway, to say nothing of
running water. We named this plateau Red Nose Point because the ground
here has a reddish color. Supposedly, every now and then a mountain lion
shows up in this area, but except for their dried droppings, we didn’t come
upon any; sadly, the farmers hunt these beautiful animals, despite the fact
that they don’t need even to protect any cattle.
I can feel the strenuousness of the last days in my bones, because I have
carried shovels and pickaxes up the mountain, and I secretly wonder why I
do this to myself in my spare time. Sure, I know our plan—we want to
excavate “fish lizards,” so-called ichthyosaurs. But you don’t have to travel
halfway around the world to do that. There are ichthyosaurs in England and
Germany, too. In Holzmaden, southeast of Stuttgart, lies one of the most
famous sites of these prehistoric animals, right next to the motorway. So
why climb steep mountains in the North American desert? Why all the
drudgery? And why are these “fish lizards” found on a mountain at all? This
is one of the most exciting mysteries of vertebrate evolution, as I will
explain below.
The rocks of the Augusta Mountains were deposited at the bottom of a
vast ocean nearly 250 million years ago, and enclosed within them lie the
remains of the first fish lizards. And although paleontology has known
about ichthyosaurs for about two hundred years, we still don’t know exactly
how they evolved. The first complete skeleton of an ichthyosaur was found
in 1811 by twelve-year-old Mary Anning in Lyme Regis, on the south coast
of England. That was thirteen years before the very first dinosaur was
scientifically described. We do know that ichthyosaurs were reptiles and
that their ancestors lived on land—but who those ancestors were is still a
mystery today.
What makes solving this mystery so difficult is the highly derived
skeleton of ichthyosaurs. The shape of their bones changed greatly during
their evolution, and as a result of their extreme adaptation to life in the sea,
they differed significantly from their land-dwelling ancestors. They no
longer possessed hands or feet, but rather finlike limbs, and their head was
long with a pointy snout, achieving the best possible aquadynamics. The
neck region was short, and the occiput was immediately followed by the
shoulder girdle. These animals were perfectly adapted to life in the open
sea, and later forms became fast, persistent swimmers due to their tail fin,
which resembled that of sharks, and their torpedo-shaped body. Eventually,
they moved only their caudal fin when swimming, while the rest of the
body remained immobile. Even the lateral fins were no longer used for
locomotion, but only for stabilization in the water and for maneuvering.
When propulsion is generated solely by a powerful stroke of the caudal fin,
this mode of locomotion is called thunniform. Sharks, dolphins, and tuna
(the term “thunniform” is actually derived from tuna) move in the same
way. However, their similar body shapes are not an indication of a close
relationship, but merely indicate similar ways of life and adaptation to the
dense medium in which they move. Living in the open ocean forced these
animals to adopt the same body shape during their evolutions because water
resistance is much stronger than air resistance on land. The evolutionary
pressure toward a streamlined shape is particularly strong in marine
animals, which is why animals in the ocean resemble each other more
closely. When completely different animals evolve toward the same body
type, it is called convergent evolution. The physique of the ichthyosaurs
was so strongly influenced by their life in the sea that it was no longer
comparable with that of their ancestors, which is why it’s difficult to
identify and find these ancestors—not least because, since they lived on
land, they do not occur in the same sedimentary rocks as their swimming
descendants.
The origin of ichthyosaurs is particularly fascinating and mysterious
because we find these animals already in rocks that are 250 million years
old. There are no finds in the Paleozoic, before the Triassic, but about a
million years after the most devastating mass extinction the world has ever
seen, the ichthyosaurs suddenly appear—abruptly and en masse! In China
and Japan, in Spitsbergen and here in Nevada, we see hundreds, maybe
even thousands of fossils of these marine reptiles in the fossil record. Where
they came from, no one really knows. But these animals are all completely
adapted to life in the sea. Now, a million years may sound like an
unimaginably long time from a human perspective, but considering the long
evolutionary history of life on our planet and the severity of the
consequences of the mass extinction at the end of the Permian, it’s no more
than the blink of an eye. And in that relatively short period of time, their
skeletons completely transformed, their hands became fins, and they
evolved from egg-laying land dwellers to viviparous deep-sea swimmers. In
the Augusta Mountains of Nevada, we find some of the oldest ichthyosaurs,
the likes of which are otherwise found only near the Chinese city of Hefei,
about three hundred miles west of Shanghai. The state of preservation of the
animals in Nevada is mostly good; bones and skeletons are even preserved
three-dimensionally. In Holzmaden, Germany, on the other hand, fossils of
the fish lizards were flattened over millions of years as a result of rock
compression (diagenesis). The rock beds in Nevada are not completely
horizontal, but dip slightly to the south. Thus, as we hike north, from one
canyon to the next, we can travel back through time, so to speak, looking at
older and older rocks and gradually approaching the Permian–Triassic
boundary. The mountain range stretches from north to south, and our
excavation takes us into two canyons: the Favret Canyon, whose rocks date
from the Middle Triassic, and the Mustang Canyon, farther north, with
sediments from the older Lower Triassic.
It’s lunchtime, the sun is blazing even hotter from the sky, and I eat my
sandwich under a large juniper bush. A small lizard squints enviously from
a stone at the sandwich. These little animals are barely larger than sand
lizards, but are related to the large iguanas. They have a sand-colored body
with brown spots and bright yellow feet. On the neck are two noticeable
black bands separated by a white stripe. They’re called Great Basin
Collared Lizards and I think they’re downright pretty. The “Great Basin” in
their name refers to the geological structure that includes the Augusta
Mountains. It is located in the Western United States and stretches from
Oregon in the north to Mexico in the south.
I drink another big sip of water from my gallon, into which I mixed blue
Gatorade powder that morning. In the mountain desert here, the air is so dry
that you don’t even notice how much you sweat, because it evaporates
immediately. So, you have to be mindful to drink enough—especially if you
do any physical labor here. Back to work. In front of me are eleven large
rocks, which are about ten feet across when lined up; together they form a
sausage-shaped concretion.

A concretion is a mineral aggregate that is often round or

spherical. In this case, it is particularly long and narrow, flat
and oval. It consists of a hard, fine-grained sediment that was
cemented by pore water and grew outward from a
crystallization center over time.

In this concretion, the crystallization center is a marine reptile that we

excavated in the days before. It is called Omphalosaurus and it is an
extremely unusual animal. The concretion measures only about ten feet in
length, but since the animal’s tail is missing, it could have been about fifty
feet long. Despite its enormous size, it was probably not a predator at the
top of the food chain but instead fed mostly on hard-shelled animals such as
the common ammonites. Ammonites are early relatives of squids, i.e.,
cephalopods, from whose shells only the head and arms protrude, as in the
Nautilus still living today. We know the eating habits of the omphalosaurs
because they had a so-called crushing dentition with which they could crush
or break open the thick shells of the ammonites. However, the form and
arrangement of the mushroom-shaped teeth still puzzle paleontologists
today—they are among the most unusual teeth in the entire animal
kingdom.
When we finally uncover the animal, we are thrilled. The fossil is better
preserved than it first appeared. In front of us lies a complete skeleton from
this group of animals, better preserved than anything that has been
excavated before. Martin Sander, one of my professors from the University
of Bonn, was at this very spot a few years earlier and saw a few small bone
fragments sticking out of the rock. Unfortunately, he didn’t have enough
time to recover the animal then and couldn’t wait to get back here. He knew
immediately what he had found there, because he is one of the leading
experts on this enigmatic group of animals and knows the anatomy of
omphalosaurs like no other.
Martin was my teacher and mentor in 2011 and supervised my thesis. He
is a great role model to me and knows the Augusta Mountains like the back
of his hand. The team also included Lars Schmitz, who was teaching at the
University of California, Davis, at the time; Martin’s PhD student Koen
Stein, who is now employed at the Free University in Brussels; Herman
Winkelhorst, a Dutch expert on marine reptiles; and me.
Now we are satisfied and lie down in turn, stretched out on the floor next
to the animal, to take photos indicating the scale of our find before the
blocks of rock are plastered. Describing this moment is not easy for me,
because even if not everyone shares my enthusiasm for ichthyosaurs or
dinosaurs, it is certainly overwhelming for anyone to stand or lie next to a
fossil that has been encased in rock for nearly a quarter of a billion years
and that they have excavated with their own hands. We are the first people
to get a glimpse of this animal. It once sank to the bottom of the sea and
was embedded in the mud even before the dinosaurs roamed the Earth.
Since then, entire continents have drifted thousands of miles apart, oceans
have opened and closed, and massive mountains have piled up and been
eroded again. Plate tectonics have caused the former ocean floor to rise and
form into mountains. These tremendous processes straighten out my view
of the world and my place in it quite profoundly. A quarter of a billion years
—that’s incredible!
However, we are not here to break records, but to try to find clues for the
solution to one of the greatest mysteries in vertebrate paleontology: Why
was there such a complex and diverse vertebrate fauna in the Lower and
Middle Triassic seas so soon after the mass extinction at the end of the
Permian, and where did these animals come from?
The sediment in which we found Omphalosaurus clearly originated from
the open sea far from shore. This is also illustrated by the many other
ichthyosaur species without crushing dentition that occur in this area. That
Omphalosaurus was also an ichthyosaur was not clear to everyone. For
example, in 2011, Ryosuke Motani, a professor of paleontology at the
University of California, Davis, expressed considerable doubt that it
belonged to the fish lizards, because Omphalosaurus was no ordinary
ichthyosaur. Finds of this animal are rare worldwide and almost always
fragmentary. Omphalosaurus probably did not have the characteristic tail
fin of its relatives but still moved sinuously through the water. Because it
also had finlike paddles, a streamlined body, and an elongated snout, it was
unable to walk on land. However, Omphalosaurus fed on hard-shelled
animals, not fish like other ichthyosaurs. For a long time, it therefore
puzzled researchers trying to determine its origin and affiliation, because
finds of this animal mostly consisted only of jaw fragments with bulbous or
spherical teeth, whose tooth enamel caps looked like the caps of
mushrooms. Characteristic skeletal elements that only occur in fish lizards,
and by which Omphalosaurus could have been clearly identified as one,
were generally absent. Thus, an assignment in the animal kingdom was
difficult. These teeth are in fact completely atypical for fish lizards. They
could even be easily confused with those of the placodonts from the
Germanic Basin. There was a shallow sea at the time of the Lower and
Middle Triassic where these lizards were common. They also had a
crushing dentition, but their teeth were arranged quite differently. But if
only parts of the skull are found and the roof of the mouth is missing, this
distinguishing feature may not be present. Examinations with a scanning
electron microscope can at most reveal differences in the enamel.
In 2011, the discussion about which animal Omphalosaurus exactly was
and in which ecosystem it occurred was in any case not yet completely
finished. Martin pleaded for an affiliation to the ichthyosaurs and a way of
life in the open sea. That day in the Augusta Mountains, the evidence for
this was right in front of us. Motani had doubted the affinity to the
ichthyosaurs and also saw gaps in Martin’s interpretation of the habitat.
This had all been difficult for Martin to prove at the time. We did know
Omphalosaurus from Nevada and Spitsbergen, which was clearly a fully
marine signal, but Omphalosaurus was absent from rocks of the same age
from the Anhui province in China. There’d been no reports of finds there,
although those sediments had a similar depositional history. Two finds from
the Germanic Basin caused us additional headache, because this was a
completely different habitat. There should not have been any omphalosaurs
there at all, because it was a shallow sea and other ichthyosaurs were
completely absent. Instead, there were placodonts, which are absent from
all other sites bearing omphalosaurs, as they probably competed with them
for food resources.
The occurrence of omphalosaurs in the shallow sea of the Germanic
Basin made little sense, and in almost two hundred years no other
omphalosaurs were ever found there except for these two fragments. Many
clues also suggest that Omphalosaurus’s diet was only found in the open
sea, far from the coast. It fed on hard-shelled organisms, all of which were
free-swimming, rather than sedentary mollusks or corals, as the placodonts
in the Germanic Basin may have eaten. Worldwide, corals recovered very
slowly from the Permian mass extinction, and it was not until about 10
million years after the Permian–Triassic transition that coral reefs had fully
regenerated. In the shallow sea, this process was somewhat faster.
 In any case, there is no evidence of coral or shell reefs in the sediments
of Nevada. The sediment consists of sea floor from the open ocean. There is
no sign of burrowing organisms, such as worms, that would have burrowed
through the substrate at the bottom of the sea. Such geobionts are also
called benthos, and burrowing, rutting, or digging at or in the ground is
known as bioturbation. In this area, however, there was no bioturbation, no
benthos, and no corals. The seabed was dead. It consisted only of sulfurous
silt with no oxygen. Instead, there were myriads of bivalves, called
daonellids and cephalopods, swimming freely in the water column.

The water column is a defined vertical section of a body of

water, extending from the surface to the bottom. It serves as a
conceptual framework to study the biological features and
processes of lakes or seas, including the presence of micro-
and macroorganisms.

In the water itself, the situation was not much better, because here, too,
there were anoxic and oxygen-deficient zones. The sediments that indicate
these zones are the so-called black shales. The daonellids and ammonites
were apparently able to cope with this oxygen-deficient environment better
than other animal species, so much so that this habitat was largely lacking
species diversity. However, the few animal species that could cope with the
adverse conditions occurred here in abundance.
The anoxic conditions may also explain why we found ten or more
ichthyosaurs during our excavation, but comparatively few fish that receive
oxygen directly from the sea through their gills. If it were lacking there,
they would not survive. Ichthyosaurs, like whales and dolphins, are not
dependent on oxygen levels in the sea and breathe atmospheric oxygen after
surfacing. Conditions for the fish lizards were less than ideal, but the low
oxygen levels in the ocean would not hinder their development as long as
they had enough to eat. So, the local story in Nevada seemed clear, even if
the mystery of the absence of Omphalosaurus in China and its supposed
presence in the Germanic Basin still remained unsolved.
Only a study by a team from Bonn led by the paleontologist Tanja
Wintrich—who, six years after our excavation in Nevada, examined the two
jaw fragments from the Germanic Basin that had been assigned to
Omphalosaurus—helped to clarify the facts. She used a scanning electron
microscope and computed tomography to visualize the complex
microstructure of the enamel and the structure of the teeth in the jaws of
these two finds. In the case of one piece from the Lower Middle Triassic
found in eastern Poland, the characteristics of the tooth-bearing fragment
provided unequivocal evidence that the jaw was indeed that of an
omphalosaur. This was indicated by the shape of the teeth, the morphology
of the enamel surface, the unique enamel microstructure, and the peculiar
tooth replacement pattern that was unique to Omphalosaurus. The teeth of
the animal are indeed amazing. They seem to erupt from the jaw almost at
random. Microcomputed tomography shows that at least two more
generations lie dormant beneath the functional teeth. However, they do not
lie directly under the erupted teeth, but are offset, with no discernible tooth
roots, and they do not grow out of sockets, but are simply stuck randomly in
the jaw. Because we do not yet fully understand this tooth structure, Tanja
always speaks of the enigma that is Omphalosaurus.
The other piece from Rüdersdorf near Berlin turned out to be a fragment
of the left upper jaw of a placodont. Here the characteristic microstructures
in the enamel were missing; the shape and the arrangement of the teeth in
the jaw were also different. Now the story made more sense. The shallow
part of the Germanic Basin was indeed devoid of omphalosaurs, while the
find in eastern Poland was located at a sea gate connecting the Germanic
Basin of the Muschelkalk with the open sea of the Tethys. The animal had
perhaps strayed or ventured too far into the basin before dying there.
The story was then rounded off when, in 2020, Martin learned that
omphalosaur teeth had very well been found in the Anhui province, and
quite a while ago, at that. Why colleagues there had not reported these finds
and why there are no scientific articles about them is difficult to understand.
But we knew nothing of all this on that Monday in the early fall of 2011.
I made a drawing of the position of the individual boulders in relation to
each other so that the preparators could rearrange them correctly again later,
back in the laboratory. Then I was allowed to wrap the blocks with
bandages dipped in plaster and tie burlap around them. This protected them
for transport. A helicopter came all the way from the town of Winnemucca
for this, to fly them down into the valley in a huge net. Koen and I flew up
to the dig site from base camp with the pilot, an elderly Vietnam veteran
named Ted. The climb, which usually took us hours, now happened in three
minutes. Ted landed the aircraft on a bare, flat spot about three hundred feet
from the omphalosaur. He stopped only until Koen and I had hopped out of
the cockpit and nimbly stooped to get far enough away under the running
rotor blades. Then we had to load the rocks into the net and attach it with a
large hook to a rope hanging down from the belly of the helicopter. While it
was approaching again from above, we wore earmuffs, yet the noise was
still considerable. We waited by the net until the helicopter was close
enough. The rotor blades were stirring up dust, and every move had to be
executed quickly and precisely so Ted could fly away promptly. It took only
a few moments to hook up the net and then away he flew with the precious
cargo. We quickly ran to a cliff and we watched the helicopter now flying
below us through the Favret Canyon toward the valley. What an impressive
sight! We followed the helicopter on foot and descended through the
canyon. Halfway down, we passed another prominent spot, which we
dubbed Beer Point. From this vantage point we could see the base camp for
the first time, and we could already anticipate the cool beer that was waiting
for us in a cooler down there. Today we had really earned it.
The concretion with the omphalosaur skeleton being recovered by helicopter. On the ground: myself
(left) and Martin Sander (right).
PHOTO BY LARS SCHMITZ

Nevada’s Ichthyosaurs: All Just One Big Family

Later, at the base camp campfire, the whole team ate dinner and drank water
and a few bottles of beer. Usually, we bought the cheapest beer brand.
Once, however, Lars spotted a local beer at the supermarket with an
ichthyosaur on its label, and he bought a six-pack of it. It was called
Ichthyosaur India Pale Ale, or Icky for short, and came from Great Basin
Brewing Co. It made us wonder why the brewery had a geological structure
in its name and offered a beer with a label that featured the state fossil. In
the United States, each state has its own official motto, flower, and animal,
and sometimes even an affiliated fossil. Nevada’s fossil is the ichthyosaur
Shonisaurus, a giant whalelike fish lizard from the Upper Triassic.
 Since our plane was taking off from Reno anyway, we decided we’d stop
by the brewery before we left. But for now we were still sitting around the
slowly dying fire talking about the day’s work and the second big
excavation we had planned for the coming days: we were going to dig up
the sea monster of the Augusta Mountains that we had discovered on the
north flank of the Favret Canyon. This monster was so huge and had such a
big mouth that it must have eaten not fish but other ichthyosaurs. At some
point, Herman cleared his throat because he felt an itch there. And when
Koen and a little later also I felt the same itch in our throats, we knew—it
was high time for our “medicine.” Lars took out a bottle of Tennessee
whiskey, and after each one of us had drunk a good slug of whiskey, the itch
immediately subsided. Solely as a prophylactic measure, Martin and Lars
also took a sip. It had become dark around us in the meantime, and the clear
sky offered a fantastic view of the Milky Way. This place was so secluded
and free of light pollution that the stars shone brighter in the sky than
anywhere else I had known before. Those evenings around the fire,
physically exhausted but satisfied with my day’s work, that untouched
nature, the camaraderie, and the significant finds make the Augusta
Mountains a very special place for me to this day.
The big monster we wanted to excavate was located in the Favret Canyon
near where we had previously recovered Omphalosaurus, about one
hundred and fifty feet farther down the steep slope. The rocks there are
from the Anisian, the lowest Middle Triassic stage, and are about 246
million years old. The round dorsal vertebrae of the animal already
weathered out of the mountain, just rolling down the slope. The dorsal
vertebrae of ichthyosaurs are very distinctive: when viewed vertically from
above, they appear circular, and in cross-section they look like an hourglass.
We suspected that this huge animal was a Thalattoarchon. This was a
massive predator, about thirty feet long and with a huge skull. Its pointed
teeth were flattened on the sides and measured at least five inches long. In
front and back they had serrated edges, with which the animal could cut
through the flesh of its prey like a steak knife when biting down.
Thalattoarchon’s full name is Thalattoarchon saurophagis, which means
“the lizard-eating ruler of the seas.” An excellent name, in my opinion.
Thalattoarchon was the very first apex predator of the seas known to
science. It could hunt extraordinarily large prey and kill them with its teeth.
Its dentition was designed so that it didn’t just swallow its prey, as most fish
do, but could bite off large chunks of flesh from it. Thalattoarchon’s teeth
were shaped in such a way that they were unsuitable for catching and eating
smaller prey. It thus depended on eating other species of fish lizards, as its
figurative name suggests. One can perhaps compare its way of life with that
of modern-day orcas. Such hunters, which can kill very large prey and are
at the top of the food chain, are also called apex predators, and their way of
life has far-reaching effects on the ecosystem. For a healthy population of
predators to be maintained, prey must be at least a hundred times more
abundant than their hunters. Statistically, there may only be about one lion
for every hundred zebras in the savannah. In the oceans, the ratio is
different from that for land animals, but animals that serve as food for
predators must still be available in sufficient numbers before the predator’s
ecological niche can even be occupied. These niches are referred to as
trophic levels.

A trophic level describes the position of an organism within

the food chain. The food chain follows a certain order, at the
beginning of which are the so-called primary producers,
which are organisms that perform photosynthesis and do not
feed on other animals. These photosynthetic organisms
include land plants and algae or lichens. They are on the first
trophic level. Herbivores, such as sheep or cows, are called
first-order consumers and are on the next trophic level. They
are followed by second-order consumers, which are
carnivores that feed on herbivores. Many ecosystems do not
go beyond this level of complexity. The second-order
consumers are then followed only by decomposers
(detritivores) such as molds or bacteria, which decompose
dead organic matter and excrement and return their nutrients
to the soil and water, making them available to plants.
 Thalattoarchon represented at least a third-order consumer—that is, a
carnivore that fed on other carnivores. Because only significantly less than
10 percent of one trophic level’s energy can be transferred to the next, there
are no ecosystems that go beyond five trophic levels. At some point it
becomes, energetically speaking, uneconomical to hunt and eat predators
because there are so few of them and the chances of a hunting success
therefore become smaller and smaller—not to mention that predators can
usually protect and defend themselves better than herbivores. Therefore, it
is absolutely amazing to find a Thalattoarchon in a world that had already
developed a complex food web shortly after the mass extinction at the end
of the Permian. That would be something like having a super lion that eats
other lions. Such things do not occur in today’s animal kingdom.
When we stood on this northern slope in 2011, shoveling away many
cubic feet of dirt and gravel with a pickaxe and a spade, we did not yet
realize how ichthyosaurs could diversify so quickly and occupy so many
ecological niches. At that moment, we only hoped to find a skull of the
animal as well preserved as possible.
The only specimen of a Thalattoarchon recovered up to that point was
missing the tip of the snout, but we hoped for better luck. We dug into the
mountain and first found the anterior part of the vertebral column, the
shoulder girdle, the humeri of the finlike forelimbs, and the occiput of the
animal, and it was soon clear that the skull was completely preserved.
However, the head, along with its snout, was still stuck horizontally in the
mountain. This meant that we had to remove about six or seven feet of
debris and overburden and work our way a good ten feet into the hill. We
dug with shovels and picks down to a thick layer of whitish-gray tuff and
then on to about eight inches above the concretion in which this animal lay
encased. At this level, there was a second thin orange layer of volcanic tuff.
This served as a useful marker for us in the field. When we saw this tuff
layer elsewhere, we knew that the rocks were of the same age and that
fossil-bearing sediments lay beneath it, increasing the likelihood of finding
more ichthyosaurs. Volcanic tuffs were also important for determining the
absolute age of rocks.
 Tuffs are igneous rocks that are ejected from the mountain
during volcanic eruptions and can spread over a large area. It
usually consists of volcanic ash and small rock fragments that
have been baked together by the extreme heat. We often find
zircons that can be used for radiometric dating enclosed
within it. If one examines the radioactive isotopes contained
within the zircons, which are subject to a natural decay
process, one can determine the age of these layers very
accurately.

Since the fossil we were excavating was directly under a layer of tuff, we
were later able to determine its age with a high degree of accuracy in the
laboratory. This was important in that other index fossils such as conodonts,
bivalves, crinoids, coralline algae, and ostracods were completely absent
here—except for the ammonites, whose biostratigraphy can also be used to
determine the age of the Triassic. Besides the ichthyosaurs, the ammonites
formed one of the groups that recovered most quickly after the Permian–
Triassic catastrophe. About one million years after “the Great Dying” they
were again abundant worldwide. It is assumed that probably only two
genera of these cephalopods survived the mass extinction at the transition to
the Triassic, but already in the lowest Triassic more than a hundred genera
can be found again. Thus, a very fine biostratigraphic classification of the
Lower Triassic is possible.

Biostratigraphy is based on the principle of fossil succession

and divides rock units with the help of fossils. According to
fossil succession, one fossil community is replaced by
another over time. Once a fossil becomes extinct and
disappears from the fossil record, it never returns in the
stratigraphy.

To our knowledge, however, there was no other record of volcanic ash in

the region at that time apart from our tuff finds. So, with these samples, we
were able to contribute radiometric data to the abundant biostratigraphic
data set for the first time. Therefore, we packed a few large bags full of
them and took them with us.
Shortly before reaching the concretion, we had to begin working more
carefully. After the thin orange layer, only brooms, brushes, screwdrivers,
and oyster knives were used. When we finally uncovered the skull, a huge
head about six feet long lay before us, completely preserved up to the tip of
the snout, just like the humerus, which measured 16.7 inches. That may not
sound very big—but for an ichthyosaur, it’s huge.
Now came the most difficult part of our work: with heavy hearts, we had
to fill up the hole again. We placed shovels and picks next to the skull,
spread a tarp over it, and carefully poured loose gravel on top of it to
protect the skeleton from weathering. We knew we would not be able to
recover the animal during this field season. The hourly rates for the
helicopter were simply too expensive and we did not have enough plaster or
a suitable crate to send the monster to Germany. We also didn’t have
enough time to get the necessary paperwork and logistics sorted. Martin
knew, however, that he would be back. We took one last longing look at this
magnificent piece, while the suspicion came over us that it might well be an
apex predator and a consumer of a third or higher order—but not a
Thalattoarchon; rather, a completely new species, instead. Indeed, this
colossus was also much larger than Thalattoarchon. We noted the GPS
coordinates and started another fossil search.
Three years later, the animal was recovered by another team, and it took
years to carve the fossil out of the rock. It wasn’t until 2021, ten years after
we first uncovered the animal’s skull, that the results were published
scientifically. It must have been about fifty-five feet long and weighed
about forty tons when it was alive. That’s almost twice as long as
Thalattoarchon and about the length and weight of a sperm whale. The
skull alone measured almost seven feet and the jaws were armed with many
pointed teeth. This monster was the first giant of the animal kingdom.
It now bears the scientific name Cymbospondylus youngorum. The genus
name means “boat vertebrae,” and the species name honors the couple Tom
and Bonda Young, who financially supported the 2014 dig. They are the
owners of Great Basin Brewing Co., which also brews the Icky beer. This
find is special because this animal is so large yet incredibly old at the same
time. While there are similarly sized animals later in the evolution of
ichthyosaurs, and some that are even larger, Cymbospondylus youngorum is
the first whale-sized marine giant in Earth’s history. This is so remarkable
because it emerged only about 2.5 million years after the very first
ichthyosaurs appeared, and its first marine ancestors were only about three
feet in size or less. This suggests an incredibly rapid evolution toward
gigantism. A similar size increase has taken nearly 25 million years for
toothed whales (odontocetes), which include orcas and the sperm whales,
ten times as long.
But the rapid rate of evolution is not the only thing that made this animal
so special. For a predator of this size to find enough food, there had to be
very stable conditions in the food web and prey had to be available in
sufficient quantities. In addition, unlike baleen whales, a tooth-bearing
animal of this size could no longer feed on small ammonites or fish because
it had to meet a much greater energy demand and was no longer nimble and
agile enough to pursue small prey. Cymbospondylus youngorum therefore
probably no longer fed on ammonites or fish—or at most on large
ammonites and very large fish—but on other, smaller ichthyosaurs. It
occupied a higher level in the trophic pyramid and was probably the top
predator in its environment. Cymbospondylus youngorum was a third-order
consumer—that is, a carnivore that ate other carnivores.
 But all this has only recently become known to paleontologists. In the fall
of 2011, we continued to search for ichthyosaurs, and Koen found a
skeleton that was given the field designation Koen 1. This is because during
excavations, finds are not initially given a scientific name because it is
impossible to be sure what genus or species they are before a detailed
examination, and because some finds represent new species of animals or
plants that do not yet have a scientific name at all. Martin found another
small Cymbospondylus, which was named Martin 1, and a second one
shortly after—Martin 2.
Lars came across a small ichthyosaur, a Phalarodon (a close relative of
the Mixosaurus), which was named Lars 1.

Lars discovered a slab with ichthyosaur ribs sticking out.

PHOTO BY LARS SCHMITZ

The following day, Martin even found a third ichthyosaur, in whose

thorax we discovered another, smaller vertebrae close to its spine, and then
two more. The large vertebrae were about two inches in diameter, the small
ones only about half an inch. They showed no signs of bite marks, and the
bone surface had not been affected by stomach acid. So the big animal had
not eaten the small ones, no—we were dealing with a pregnant female. This
caused a small sensation! The animal, which we did not call Martin 3, but
Martina, was not only a new species, but also the world’s oldest record of a
viviparous fish lizard or a viviparous reptile at that time. Prior to this
female, only two valid species of Cymbospondylus from the Augusta
Mountains were known: Cymbospondylus nichollsi, which had a body
length of about twenty-three feet, and Cymbospondylus petrinus, which
grew to thirty feet in length. Martina must have been about fourteen feet
long; later studies of her skeleton showed that she’d been fully grown and
sexually mature. Although she was smaller than the already known
cymbospondylids, her teeth were just as long, clearly identifying her as a
new species.
We spent a lot of time searching the mountainside for bone fragments and
vertebrae, which we carefully collected. We used superglue to fill in the
cracks and stabilize the remaining articulated pieces. We noted the
coordinates of each site and collected any loose pieces. Martin and I worked
alone that day because Koen, Lars, and Herman had gone into town to take
care of the shipping of the omphalosaur. We covered this site also so that
the rest of the skeleton, which was still trapped in the bedrock, was
protected and could be excavated during a future excavation. That, too,
would be three years in coming. In the end, I shouldered the loose
fragments from Martina, and we headed back down to the valley.
One reason Martin had taken me on this dig was my hiking experience
and what he considered my great resilience. I usually carried the heavy
equipment up the mountain and our fossil finds back down. I may not be the
most athletic person, but I’m used to hiking, and I’m tenacious and
downright stubborn. Even though the equipment was very heavy, I refused
to give up. A few months earlier, I had been on an excursion with Martin
and other students to the Bletterbach gorge in South Tyrol. There we had
found the fossil footprint casts of a parareptile called Pachypes dolomiticus
in the riverbed. Such casts are natural infillings, formed when footprints are
permeated with harder material that is resistant to weathering. The name of
the animal means “heavy foot from the Dolomites.” It lived immediately
before the Permian–Triassic mass extinction event and was one of the
largest herbivores at that time. The casts were quite heavy. But since I
already knew that Martin would be traveling to Nevada a little later, I was
eager to prove to him that I could handle the arduous marches in Nevada
even with heavy baggage. So, I carried the fossil footprint all the way up to
the museum, where we handed it over to the curator. I was eager to go with
him to Nevada and wanted to show Martin that I was indispensable. None
of the other students made such a formidable carrier as I did. Martin
actually took me with him to Nevada. We sometimes joked that I was a kind
of personal Sherpa for him, but I really wanted to be on the expedition to
the Augusta Mountains and was happy to take on all the hardships. So, I
carried plaster bags up the mountain and parts of the pregnant female
ichthyosaur back down. In return, I was an eyewitness to the discovery of
these spectacular finds, saw tarantulas and rattlesnakes, and experienced
alpenglow in the Nevada mountains. Once, after a change in the weather
where temperatures dropped rapidly, it even snowed on Cain Mountain, and
Lars, Martin, and I climbed the snow-covered rise. What a magnificent
spectacle of nature!
On one of the last days of our fieldwork, we went even further back into
the Earth’s past in search of ichthyosaurs in the Mustang Canyon, a little
farther north. Martin had never taken on this terrain before because there
was just so much to do in the Favret Canyon and never enough time to do it.
The Mustang Canyon was composed of Lower Triassic sediments that were
even older, going back almost to the Permian–Triassic boundary. If we
found ichthyosaurs here, they would surely be the oldest representatives of
the group. And indeed—right at the entrance of the canyon—we discovered
a place where isolated vertebrae of an ichthyosaur were weathering out of
the ground. To our great chagrin, it looked like a fossil hunter had looted
this spot and removed the skull. The Augusta Mountains are under the
jurisdiction of the Bureau of Land Management, or BLM, an agency within
the Department of the Interior, and you can’t dig here or take anything
without a permit. We possessed such a permit because our excavation was
for scientific purposes. All of our fossils, after being prepared and fully
examined, were brought back to the United States and given to a museum
collection. Fossil poachers, on the other hand, looted sites and then sold the
skeletons privately to the highest bidder. How severe the damage to the
scientific community is, when fossils are taken out of their geological
context and cannot be professionally studied, can only be adumbrated.
Whoever had stolen this fossil had prevented the discovery and description
of an unknown species.
I think it was Koen who first found the remaining pieces. His sleuthing
was remarkable, but we didn’t assign a dig number or nickname this time
because the sparse remains weren’t distinctive or diagnostic enough to
make sense of. Our irritation was understandably great. But the carelessly
left fragments were followed by Koen 2—a small, articulated ichthyosaur—
and Koen 3, also an ichthyosaur, which actually contained jaws and teeth.
These were the first ichthyosaurs from the Lower Triassic discovered in
Nevada, and they appeared to be the oldest ichthyosaurs worldwide!
In 2014, however, two scientific articles were published by the
aforementioned Japanese colleague, Ryosuke Motani, that changed the
status quo of the Nevada findings. In one of these articles, he provided the
oldest evidence of viviparity, and in the other paper he reported an
ichthyosaur find from the city of Chaohu in China that was even older than
the Mustang Canyon specimens. This toppled both Martina and Koen 2
from their thrones.
After having discovered nothing except seashells and ammonites during
the whole expedition, I finally found my first ichthyosaur in Mustang
Canyon: Armin 1. It was another small, articulated specimen that had been
deposited in the same strata as Koen 2 and Koen 3. We documented the
finds in our logbook, protected them with a cap of plaster and noted the
coordinates. Then we carefully covered them with a few shovels of soil.
There was no time left for their recovery because we had to leave the camp
the next day. To this day, these three specimens have not been
comprehensively scientifically examined. Although we always have to
expect that poachers could steal the pieces, the Augusta Mountains are so
remote and the area is so vast that it is difficult to find the excavation sites
without GPS data.
Our campaign was immensely successful. This was partly due to the
experience and sleuthing of the participants, but also because the fossils
there are particularly abundant. This is related to the environmental
conditions of the Triassic, where dead animals that sank to the seafloor were
not eaten by scavengers because of the anoxic conditions there, but instead
quickly covered by sediment.
Much of what we discovered during this and subsequent excavations
could not be accurately interpreted at the time, and our records were
eventually surpassed. Science just takes its time, and records themselves are
ultimately unimportant because paleontology is about understanding nature
and evolution.
Searching for and recovering fossils in the field is tedious and costly, but
the biggest cost in the end is the preparation in the lab, which also takes the
most time. Our dig took place many years ago, but only now are
publications about these finds coming out. An article on the pregnant
female ichthyosaur came out in 2020 and one on the Augusta Mountains
monster in late 2021. The fact that these publications on our Nevada finds
have been so long in coming is largely related to the preparation of these
animals. It is extremely difficult and incredibly time-consuming because the
ichthyosaurs were stuck in concretions. As mentioned earlier, these
aggregates are formed by circulating pore waters sintering out from a
crystallized center within an otherwise fine-grained sediment. This makes
the nodule very heavy, solid, and particularly hard. The fossilized bones and
the surrounding rock matrix of the concretion have almost the same density
and cannot be easily separated along the divisional surface between bone
and matrix like with many other fossils.
The Omphalosaurus was particularly difficult to prepare. Several
preparators in Germany and the United States have worked on it in vain.
But although the preparation is not yet finished, we can already say a lot
about the animal. Until our find in Nevada, no complete skeleton or even
nearly complete partial skeleton of an Omphalosaurus had been found and
scientifically described anywhere in the world. Our find, however,
contained almost all of the dorsal vertebrae, so today there is no doubt that
Omphalosaurus clearly belonged to the ichthyosaurian clade.
The pregnant female was just as difficult to prepare, and it is only thanks
to the excellent work of the preparator Olaf Dülfer at the University of
Bonn’s Institute of Geosciences that we can now admire this animal, freed
from its stone prison. It takes skill and a great deal of experience to prepare
such a fossil so that it can be examined in detail. Because of this special
achievement, the animal was finally named Cymbospondylus duelferi in
honor of the preparator.
The exact phylogenetic analysis of all ichthyosaurs from the Augusta
Mountains ultimately showed that they had all been closely related and
were basically only larger or smaller Cymbospondylus. So, all the pieces of
the puzzle finally fit together here, too: Cymbospondylus was apparently
one of those ichthyosaurs that successfully occupied the predator’s niche in
the Middle Triassic seas because they had no competition from other animal
groups. The descendants of this generalist rapidly diversified and colonized
different habitats, where large, medium, and small forms fed on different
prey and rapidly split into new species. Because no other animals competed
with it, Cymbospondylus was able to produce specialists in a wide variety of
niches.
So, the pregnant fish lizard lady was a new species of the
cymbospondylian genus (Cymbospondylus duelferi), and the giant monster
turned out to be a Cymbospondylus of another new species
(Cymbospondylus youngorum). Their close relationship explains why the
ichthyosaurian fauna in Nevada was able to diversify so much in such a
short time, and how a highly complex food web was able to reemerge so
quickly after the mass extinction.
The smaller ichthyosaurs from Mustang Canyon have not yet been fully
prepared, but it will be exciting to find out if they are related to the
cymbospondylids. This would bring us a little closer to the origins of this
species-rich family. It is almost certain that they are previously unknown
species. Whether they all belonged to the same or different species will
soon become clear. With their exploration, we are moving one step closer to
the mysterious origin of the ichthyosaurs.

Of Strange Monsters and Underwater Flight

In Europe, in the Germanic Basin, the underwater world of the Middle

Triassic looked quite different. Here reefs recovered from the mass
extinction somewhat faster, and the continental shelves were soon full of
life again. The strangest organisms bustled about, both on the beaches and
in the water. Today, remains of these unusual creatures can be found on the
German-Dutch border in a quarry near Winterswijk. There are sediments
containing land creatures that came to the beach at low tide and went in
search of food. Their visits were so numerous that the slabs with their tracks
are often called “the dance floor.” We find traces of an extinct group of
herbivorous reptiles called rhynchosaurs, and of chirotheria, the ancestors
of crocodiles, whose five-toed tracks look like human handprints, which is
where their name comes from (chirotherium, “the hand animal”). In this
quarry, there are also footprints associated with another group of crocodiles,
the aetosaurs. My favorite track from Winterswijk is called
Procolophonichnium. It belongs to an extinct group of reptiles that have
scales on the soles of their feet. The limestone in which they are preserved
is so fine-grained that even such delicate structures have been preserved. If
you take a flashlight in dim light and hold it close to the surface at an acute
angle, you can produce grazing light and see the delicate scales particularly
well. Because the deposits in Winterswijk are of both terrestrial and marine
origin, we find countless footprints from land animals, though no bones; it’s
a different story for marine creatures. In the sea, we see much more unusual
animals, which appear in the fossil record only for a short time during the
Triassic and then disappear again. There are, for example, small marine
archosaurs called helveticosaurs, or the placodonts (“tablet teeth”), which,
like Omphalosaurus, had a crushing dentition, but stayed in coastal regions
and fed on sessile hard-shelled animals. Other groups included the
nothosaurs and—one of the strangest animals of all—Tanystropheus. It had
a greatly elongated neck, which was as long as its torso and tail combined
but consisted of only twelve or thirteen vertebrae.
All these animals are also found in sediments of the same age on Monte
San Giorgio. This mountain is located on the border between Switzerland
and Italy, and there is a phenomenal view over Lake Lugano from its
summit. On Monte San Giorgio, you can find, among others, the
pachypleurosaurs, or “thick-ribbed lizards.” They belong to the group of
“lizard flippers” (Sauropterygia), which were still small in the Middle
Triassic, but soon grew larger and larger, rivaling the ichthyosaurs. The
most successful group within the lizard flippers was the plesiosaurs. They
were extremely diverse and one of the longest-lived marine reptile groups.
Plesiosaurs had a unique and enigmatic body shape that we do not see in the
animal kingdom today. They possessed four pointed, evenly shaped fins, a
long, stiffened neck, and a small head. For nearly two centuries, plesiosaurs
were thought to have appeared only after the end-Triassic mass extinction
in the Jurassic, until skeletal remains belonging to a Triassic plesiosaur
were found in Bonenburg near Paderborn, Germany. The animal was named
Rhaeticosaurus, “the lizard from the Rhaetian”—the Rhaetian being the
uppermost stage of the Upper Triassic, from about 208.5 to 201.3 million
years ago. Histological studies indicate rapid growth and increased
metabolic rates in plesiosaurs, which are interpreted as an adaptation to
swimming and hunting in the open ocean. The existence of Rhaeticosaurus
supports the hypothesis that an open-sea lifestyle facilitated plesiosaur
survival beyond the Triassic.
The extraordinary anatomy of plesiosaurs, combined with their active
metabolism, enabled them to use a mode of locomotion unique in the
animal kingdom: they were able to use their four uniform fins like wings for
what is known as underwater flight. No other group of animals is capable of
this. Today, penguins also move forward underwater in this way, but they
only use their two wings; their legs and feet are not involved.

Mounted plesiosaurid Cryptoclidus in Bonn, Germany, with its flippers in underwater flight posture.
PHOTO BY THE AUTHOR

Whether plesiosaurs really used their fins for underwater flight or for
rowing was not clear for a long time. To answer this question, it was not
enough to reconstruct only the arm and leg muscles; the pelvic muscles and
the muscles of the phalanges that form the winglike fins also had to be
considered. Hydrodynamics studies show that muscles serving fin rotation
in particular are necessary for efficient underwater flight. To reconstruct
muscle groups of an extinct species, we must first look closely at the
muscle attachment grooves on their bones and compare them with those of
closely related groups still living today, such as the turtles, crocodiles, and
squamates (scaled reptiles such as lizards and snakes). We also need to
understand the corresponding muscle functions in sea turtles, penguins, sea
lions, and whales to draw reliable conclusions about how the relevant
muscle groups work in secondarily aquatic animals. Studies show that
plesiosaurs were very capable of wing flapping with the front and rear
extremities, while other muscles were able to rotate the front and rear fins
on their longitudinal axes during upward and downward strokes.

OceanofPDF.com
OceanofPDF.com
 Chapter 2

New Life on Land

Survival in Hell: The Road to Dinosaur Domination

On land, Earth during the Lower Triassic was characterized by terrible
droughts, wildfires, and a global average temperature of an unbearable
113°F. At this time, all landmasses were still united into a single
supercontinent before they drifted farther and farther apart over many
millions of years. This primeval landmass is called Pangaea. While there
was still isolated vegetation on the coasts or in the mountains, enormous
deserts larger than the Sahara spread out across the interior of Pangaea.
Despite all of these ecological difficulties, one of the most beautiful
fossils I know of comes from this very period of the Earth’s history. My
colleague Vincent Fernandez and his team found it in Africa in 2013, then
studied and scientifically described it at the multinational research facility
ESRF—the European Synchrotron Radiation Facility in Grenoble—using
microcomputed tomography. The fossil is not only exceptionally beautiful,
but also gives us unique insight into the harsh living conditions around 250
million years ago, as well as the strategies animals used to cope with them.
It helps decipher the mystery of how some animals survived at all in this
harsh, absolutely hostile, downright infernal environment, and why life on
Earth was able to persist after this catastrophe. I met Vincent Fernandez at
the ESRF while we were doing research together on a project about the
evolution of modern bony fish, scanning ribs and jaws from about a
hundred species. Vincent’s 2013 paper, however, is not about fish, but about
said fossil, which shows two animals of different genera. They lie in a
burrow, nestled close together, united in death. Exploring the circumstances
of how these two different animals died in the same burrow reminds me of a
Sherlock Holmes mystery, where meticulous detective work, tiny clues, and
clever deductions lead to the solving of a complicated case.
In the harsh environment at the beginning of the Triassic, every day was
a struggle for survival. Today, rocks that can tell us about this drama are
found in, among other places, the Main Karoo Basin in South Africa, the
stratigraphic sequence of which covers a period from before the Permian to
the Jurassic. To understand what happened shortly after the Permian mass
extinction, we have to look at the rocks from the Lystrosaurus Assemblage
Zone, a small section of fossil-rich strata within the Karoo rock sequence
that were deposited between 251 and 249 million years ago, immediately
after the mass extinction event. Vincent’s spectacular fossil came from this
very zone. In these earliest layers of the South African Triassic, one finds
many fossilized bones, but also countless burrows and subterranean
structures, which one might imagine as oversized mole tunnels. They are,
however, no longer hollow, and are now filled with sediment.
A burrowing lifestyle is a common adaptation in mammals today. It is
useful in raising young and serves as protection from predators and extreme
weather conditions. In the strata of the Karoo Basin, we find hundreds of
such so-called ichnofossils immediately after the Permian–Triassic
boundary.

The footprints, skin impressions, tiny boreholes, and burrow

structures of extinct animals are called trace fossils or
ichnofossils.

The large number of burial structures in South Africa suggests that a

burrowing lifestyle was widespread among numerous tetrapods more than
250 million years ago.

Tetrapods are all land-living vertebrates, i.e., amphibians,

reptiles, mammals, and birds.
 The period from which these fossils date is characterized by the harsh
climatic conditions of the Karoo, and the disproportionate density of
ichnofossils reflects the crucial survival strategy of many tetrapods during
this time. Sometimes cave fillings still contain the remains of their creators,
providing us with important information about the vertebrate ecology of the
time. Despite the abundance of burrow casts, however, documented finds of
burrowers within them remain rare, obscuring the burrowing lifestyle of
many prehistoric animals. From this we can theorize that burrows provided
a stable and secure environment for animals of the earliest Triassic. There is
evidence that points to seasonal periods of dormancy for their inhabitants,
probably caused by resource scarcity from things like prolonged droughts.
The variety of cave forms suggests that these were retreats for different
animals and were excavated for different purposes. The most common
producers of these trace fossils were so-called therapsids. These are
amniotes, from which today’s mammals evolved.

Amniotes are a large group of vertebrates that includes all

terrestrial vertebrates, with the exception of amphibians.
Representatives of the amniotes possess the ability to
reproduce completely independently from water. Their
development occurs in an amniotic sac filled with amniotic
fluid, rather than through a fully aquatic tadpole stage.
Amniotes include reptiles, birds, and mammals. Humans are
amniotes, too.

Therapsids probably originated in the early Permian and, excluding

present-day mammals, had their heyday in the Permian and early Triassic,
where they were the dominant predators in some ecosystems. Then the first
dinosaurs appeared, gradually replacing them at the top of the food chain.
The therapsids were menacing predators of considerable size, but they were
relegated to a niche existence by the dinosaurs from the Late Triassic
onward. After that, for about 150 million years, there were only small to
rodent-sized nocturnal creatures that ventured out of the thicket only after
sunset, when the big dinosaurs were asleep.
But at the beginning of the Triassic, these animals were larger—and
predatory. One of them was Thrinaxodon. It belongs to the subgroup of
cynodonts, or “dog-teeth,” named for their canine teeth, which are similar
to those of dogs. They first appear in the fossil record as early as 270
million years ago. At that time, the world was still suited to them. Plants
were still growing lushly and providing food for their prey, but as a result of
the end-Permian cataclysm, the vegetation was damaged in many places.
Wildfires, air pollution, and acid rain destroyed the lush forests, causing
atmospheric oxygen levels to drop from about 30 to around 10 to 15
percent. The surviving cynodonts now found themselves in an entirely new
world.
Thrinaxodon has now been identified as the main producer of these
burrows in South Africa. Sometimes one of these animals lay curled up in
its burrow. Thrinaxodon was a carnivore with sharp teeth and grew to be
about two feet long. Its legs were under its body and it had a rather short
tail. Small depressions were discovered on the sides of Thrinaxodon’s skull,
indicating whiskers, but were not necessarily evidence of full-body fur. The
whiskers probably helped the animal orient itself in the darkness of its
burrows. Its burrowing lifestyle is interpreted as an adaptation to extremely
hot summers. Studies of the bone microstructure of cynodonts indicate that
they spent extended periods of time resting in their burrows. Today, we
know such resting phases from, for example, bears, which hibernate. But
the cynodonts made their burrows for resting during the summer. This
aestivating behavior in therapsids is a survival strategy during the extremely
hot summer months.
 Aestivation is a form of summer hibernation in which the
metabolism is slowed down, and all activity is suspended or
shut down altogether as an adaptation to adverse
environmental conditions. It can occur during periods of heat
or drought, and under arid climates it can occur seasonally.
Estivating behavior still exists today in some moth and snail
species.

After Vincent Fernandez examined the unprepared fossil mentioned

earlier using synchrotron imaging, he knew that the rock block contained
Thrinaxodon (a few parts of the skeleton even peeked out at the surface),
but remarkably, the data also showed a second skeleton. It was even more
of a surprise when it became clear that this was a completely different
species! Vincent and his team had discovered a unique association of two
complete skeletons, lying side by side: a cynodont Thrinaxodon and an
amphibian called Broomistega, embedded in the same sandstone grave.
Here, one must think of the Broomistega as an oversized, potbellied
salamander. Why the two animals were in the same burrow together was
completely unclear at first. But, for the first time, this material provided the
opportunity to study behavioral interactions between two unrelated Triassic
animal species. The team wanted to determine if this interaction was
random or if it offered advantages to one of the animals involved.
Synchrotron data showed that the amphibian, a juvenile, had had seven
broken ribs but had healed. The healing process provided evidence that the
trauma had occurred several weeks before its death and had not been caused
by the Thrinaxodon. The fractures had not been fatal, but they had certainly
severely affected the animal’s locomotion and respiration. Both skeletons
are exquisitely preserved, in complete anatomical articulation and in natural
posture, suggesting that the animals were buried with soft tissues intact,
including skin and muscle, and possibly even alive. Broomistega was not
capable of digging a hole in the ground, and therefore may not have been
the original burrow dweller, but only a guest. Amphibians such as
Broomistega are well adapted to life in water and therefore do not have
bones as sturdy as land dwellers. Their body structure was unsuitable to
build burrows by themselves. Although there is no clear anatomical
evidence that Thrinaxodon was an active burrower, it can be assumed, since
several animals of this species have already been found in burrows. Thus,
Thrinaxodon was probably the primary occupant and probably the builder
of the small burrow, and Broomistega only a visitor. Because the amphibian
and the cynodont lie closely entwined in this burrow, it is obvious that they
died at the same time and that Broomistega was not washed into the hole
afterward by chance. Otherwise, the animals would lie behind each other.
A prey-predator relationship is also unlikely, as both skeletons are almost
completely intact and neither animal gnawed or bit the other. The
Broomistega, in addition to its rib fractures, has a bite mark over its left eye
that could have been caused by the canine teeth of a cynodont. But the
distance between the two holes in its bone is 16.5 millimeters, while the
canines of the Thrinaxodon in the burrow, however, are only thirteen
millimeters apart. So, this cynodont was not the culprit. The Broomistega
seems to have escaped from an even larger animal before that. Perhaps it
was from this encounter that it had its rib fractures, too. The poor animal
obviously had a rough life.
Another consideration was that the Thrinaxodon may have dragged the
large salamander into its burrow to eat it later. Storing food in burrows does
occur in the animal kingdom, but it usually does not involve hoarding
perishable food. Keeping live animals as food in burrows is rather unusual,
especially in hot climates that favor rapid decomposition of the carcass.
Given the absence of injuries attributable to this Thrinaxodon and the low
probability of it hoarding perishable food, the above hypothesis can be
safely ruled out. Vertebrates of different genera occur together in burrows in
the present animal kingdom only when external factors such as foraging,
predator avoidance, or harsh climatic conditions play a role. For example,
when one of the two species leaves the burrow, an intruder may chase away
the host, or when the host tolerates the presence of the new guest.
Especially in harsh climates, there are still cases among terrestrial animals
today where guests are tolerated by a host. Such communal living offers
advantages in warding off predators and also increases the likelihood of
survival of the animals involved through shared vigilance. Nevertheless,
such cases remain the exception. But juvenile amphibians alive today have
been known to temporarily use burrows as protective retreats. This could
also be a plausible explanation for the presence of the Broomistega juvenile.
The fact that the Broomistega was not attacked or injured by the presumed
Thrinaxodon host also suggests that the cynodont tolerated the intruder’s
presence or was unable to chase it away. Perhaps the Thrinaxodon was
already dead or paralyzed. However, if it had been dead before its burrow
was flooded, rigor mortis probably would have set in long ago. But the
lateral curvature of its spine indicates a mortal agony shortly before
drowning in the mud. Thus, Thrinaxodon either died shortly before the
flooding, and rigor mortis had already released, or its death was related to
the flooding. Since the assumption that Thrinaxodon died before the flood
requires exceptional circumstances, it seems more plausible that the
cynodont tolerated the salamander in its burrow. The time of Broomistega’s
death can be reasonably correlated to the flood and argues for voluntary
entry into the burrow. Its curved vertebral posture also suggests that it died
shortly before the flood event or during the flood.
In the Early Triassic, the climate was dry, and the drought was
interrupted only once a year by a violent monsoon season. During the
hottest months the animals became immobile, which is why aestivation can
also be used as an explanation for the therapsid skeletons in the Karoo
Basin burrows. In particular, the mostly dormant or curled-up position of
the animal skeletons supports this assumption. Studies of aestivation in
contemporary mammals indicate that prolonged inactivity and fasting can
lead to growth arrest, particularly in the arm and leg bones. Arrested growth
or growth interruptions can be detected in bone cross-sections by various
marks and lines.

Lines of arrested growth are referred to as LAGs. Such LAGs

can be found in ectothermic animals, but normally they do
not occur in mammals. However, LAGs have already been
observed in the long bones of several therapsids, some of
which were discovered in curled-up positions, indicating their
estivating behavior.

Interestingly, however, bone-histological studies of Thrinaxodon show no

such LAGs. We must therefore assume that a burrowing lifestyle provided a
continuous growth rate for the animal. Burrows mitigated extreme heat, and
strong climatic fluctuations were balanced by stable ambient temperatures
and constant humidity inside them. This helped these animals regulate their
body temperature under extreme climatic conditions, and they were also
able to cope more easily with the lack of resources. The Thrinaxodon was
able to survive without summer hibernation in a torpor stage.

Torpor describes a sleep state that occurs in small mammals

or birds and is comparable to a kind of lethargy in which
metabolic processes are slowed down to a minimum. The
body is kept “on the back burner.” The animals usually
remain in torpor in a state of body rigidity. While in this state,
they can survive for a long time without water or food.

Torpor, which can be self-initiated by mammals under extreme weather

conditions, consists—in contrast to an uninterrupted estivation—of several
short periods of rest lasting only for a few days each. Lines of arrested
growth become more indistinct or are completely absent during this period,
so that they can no longer be detected in the cross-sections of arm and leg
bones. Since the growth of the animal does not stop completely, the
characteristic LAGs are missing.
The unusual residential community of Thrinaxodon and Broomistega,
and also the numerous other Thrinaxodon specimens in curled-up body
postures, suggest that this cynodont genus retreated into its burrow for
periods of dormancy. However, in the absence of histomorphological
markers indicating an interruption of bone growth, these periods seem to
have been short-lived. Such temporary periods of torpor are a primitive
feature in mammals and thus more likely than the specialized metabolism of
hibernation. In this mammalian ancestor, a certain plasticity in metabolism
still existed, meaning that these animals could slow it down when external
conditions became unfavorable. Humans are not able to do this. If we do
not have enough food or water for a long period of time, we die. The ability
to escape threatening climatic conditions via a burrowing lifestyle, and thus
to survive a food shortage longer, certainly contributed to the survival of
small-and medium-sized cynodont animals in the Permian–Triassic crisis,
as did the variability or plasticity of their metabolism.
But when the golden era of the big amphibians and cynodonts was over,
small nimble bipedal lizards were already running around the burrow of the
odd couple who’d been united in death. These bipedal lizards coped much
better with the extreme temperatures. And so, the death of Thrinaxodon and
Broomistega is also symbolic of the demise of their respective animal
clades. The ordeal of these unequal co-inhabitants fatefully casts its shadow
ahead and also heralds the demise of their closest relatives.

The Dragon Who Wanted to Be a Tyrannosaurus

The small bipedal lizards that roamed around the Thrinaxodon burrow were
the archosaurs. Archosaurs consist of, simply put, all crocodile relatives and
dinosaurs.
If archosaurs are considered the last common ancestors of crocodilians
and birds and all their descendants, then this group originated in the earliest
Lower Triassic. Other definitions do not refer to a pure relationship, but to
characteristics in the skeleton that are unique to this group.

Such exclusive characteristics within a group are called

synapomorphies. Archosaurian synapomorphies include teeth
set in sockets deeply rooted in the jaw, a large process on the
shaft on the posterior side of the femur, at least two lumbar
vertebrae fused with the pelvic bones, a cranial opening in
front of the eye and two behind it, and an opening in the
lower jaw.

If one follows this definition, one can find the first representatives
already in the Kupferschiefer (German for “Copper Shale”) sedimentary
unit from the uppermost Permian, which is about 256 to 259 million years
old. Such anatomical features seem unimpressive at first sight, but they
made the archosaurs the real superheroes of the Triassic—or possibly, as
we’ll discuss later, the supervillains.
The cranial opening in front of the orbit (the so-called antorbital window)
reduced the weight of the skull, which was relatively large in early
archosaurs, similar to modern crocodiles. The mandibular windows
probably also reduced the weight of the jaw in some forms and better
distributed the bite forces, allowing these animals to bite harder and more
efficiently. The large process on the shaft on the posterior side of the femur
provided a large surface for muscle attachments. Stronger muscles enabled
early archosaurs to walk upright and helped them or their immediate
ancestors survive the catastrophic Permian–Triassic extinction event. The
upright gait also made it easier for archosaurs to breathe while moving
around. It freed them from the so-called Carrier’s constraint found in
lizards.

Carrier’s constraint is a phenomenon where air-breathing

vertebrates with two lungs must flex their bodies laterally
during locomotion and have difficulty breathing while
walking. The sideways flexing expands one lung and
compresses the other, forcing the used air from one side of
the lung into the other and preventing it from being fully
exhaled to make room for fresh air. Lizards, therefore, can
only do short sprints and must rest afterwards to breathe
deeply.

In the Late Triassic, animals with Carrier’s constraint were hunted by

bipedal species that did not have this disadvantage. Breathing was
especially important in the Triassic because the oxygen content in the air
had dropped drastically since the Permian and was only slowly rising again.
In this context, too, the archosaurs had a decisive advantage: they possessed
birdlike lungs! There are a few important differences in the way lung
respiration works in birds compared to other vertebrates:
 Bird lungs maintain a constant volume during respiration.
They are ventilated unidirectionally, which means that air flows
only in one direction at all times.

Contrary to that, in human lungs, breathing air moves back and forth,
bidirectionally, as if in a tidal flow. This rhythm is the same in all mammals,
with the lungs inflated and compressed with the help of the diaphragm,
almost like a bellows. However, to achieve unidirectional flow, various air
sacs in the body are inflated and deflated in a complex sequence, like a
whole series of interconnected bellows. The lungs, located midway between
the air sacs, are supplied with fresh air in one direction during both
inhalation and exhalation. The air sacs fill with this fresh air by expanding
the chest and the abdominal cavity. The sternum swings forward and
downward while the ribs and chest wall move laterally. Exhalation is
caused by compression of the air sacs with the help of the skeletal
musculature. In a world with little atmospheric oxygen, this was a crucial
advantage.
Now unidirectional lungs have also been observed in monitor lizards and
iguanas. This could mean that this type of lung evolved before the
archosaurs split off from the other lizards. If true, this lung design would
have formed as early as the Permian, but there is no fossil evidence for this
so far. But it could also mean that this concept evolved several times and
independently, i.e., convergently, in the animal kingdom.
Without the bronchial system within the lung tissue, however, iguanas
lack a key innovation that enhances the airflow pattern in birds enough to
enable them to engage in active flight. This bronchial system was likely
favored by natural selection because it facilitated metabolic processes
during strenuous activity. And while the unidirectional lung may have
evolved as early as the Permian, we don’t see this particular bronchial
system in the lungs until after the divergence of crocodilians and dinosaurs
in the Triassic, which correlates beautifully with oxygen scarcity in the
atmosphere. A super-lung, while the other animals were running out of air
—this brought the development of the dinosaurs furiously forward and
displaced more and more other animals from their habitats.
But not everything was about breathing. To be a successful predator, you
also had to have big jaws with strong teeth and be strong enough to kill
your prey. So sometimes sheer muscle helped. But if you wanted to hunt
large prey, you also had to be able to chomp them into bite-sized pieces
after killing them. This meant cutting through bones to separate body parts
from the carcass.
The ability to crush or even consume bones as a food source is called
osteophagy. Osteophagic feeding behavior, however, seems to have been
rare in carnivorous dinosaurs. This can be well explained if we remember
that dinosaur skulls were very lightly built. They had one cranial opening in
front of the eye and two behind it, and an opening on the lower jaw to
optimally distribute and relieve stresses generated during biting. All
dinosaurs possessed these openings; they are synapomorphies of
archosaurs. In addition, the teeth of predatory dinosaurs were narrow and
bladelike. They could cut meat very well with them, but they could not
crush bones. These teeth would have usually broken off or would have
chipped if the animals had attempted an osteophagic diet. Therefore, we
rarely find bite marks on bones in ecosystems dominated by dinosaurs.

The tyrannosaurids of the Late Cretaceous are a major exception. They

had massive skulls and thick, robust teeth rooted deeply in their jaws. In
these animals, we often see worn teeth in their jaws and deep bite marks on
the bones of their prey that match with the shape of the teeth of
Tyrannosaurus. In the absence of such bite marks, it is difficult for us to say
with certainty what the predators ate. Of course, we can use the fossil
record to determine which animals might have likely been prey, but that is
very imprecise. Here, the fossilized stomach contents of dinosaurs prove
much more useful. Unfortunately, fossils that reveal their last meal are
relatively rare. In contrast, it is much more common to find the fossilized
feces of dinosaurs, the so-called coprolites, through which conclusions can
be drawn about their eating habits. The coprolites of carnivores are found
much more frequently than those of herbivores because they contain
phosphate and therefore have much greater preservation potential.
However, in some birds, clues about their diet can be found without having
to examine their feces: owls and other birds of prey regurgitate hard-to-
digest remains of their prey. These are called spit balls or pellets, which can
also fossilize and are then called regurgialites.
One colleague who knows about the coprolites of archosaurs like no
other is Martin Qvarnström of Uppsala University in Sweden. I met him at
an academic conference in Albuquerque, New Mexico, where he gave a talk
about a paper on Upper Triassic coprolites in Poland. He had scanned these
coprolites at the ESRF using synchrotron microtomography, and his talk in
Albuquerque was impressive. Martin has a very casual yet confident way of
talking about his field; he is able to explain complex issues in a simple way
and to captivate the audience.
Martin examined bone-rich coprolites and regurgialites from an
archosaur whose anatomy was already very similar to that of the dinosaurs.
They were found from the latest Middle Triassic and the early Upper
Triassic in Poland. This archosaur bears the scientific name Smok wawelski.
The animal was named after the dragon of Wawel from the Polish folk tale.
So, Smok has nothing to do with the dragon Smaug who watches over a
treasure of gold in a mountain from Tolkien’s novel The Hobbit. This is not
a character from a fantasy novel, but a real predator from the Triassic.
The peculiar skull structure of the animal and the contents of its
coprolites suggest that it was able to crush and consume bone. Some other
anatomical features also suggest osteophagy in Smok. For example, the
Wawel dragon had a massive head and robust body like the tyrannosaurs.
Smok is thus visually similar to Tyrannosaurus, providing evidence of
convergent evolution associated with a similar feeding ecology.
The composition of the coprolites of Smok confirm what the skull
structure already suggests. Half of their content consists of tooth and bone
fragments. However, the tooth fragments are not from its prey, but rather
broken tips of its own teeth, which it swallowed with its food. It ate a rich
diet consisting of fish, small mammals and lizards, giant dicynodontians,
and large temnospondyls. The dicynodontians and temnospondyls were so
large that Smok could not swallow them all at once, and many of the
salamander-like creatures had bony skin plates that it had to bite through
first.
Another animal from Poland was even more similar to dinosaurs:
Silesaurus. It belongs to a group that we call the Dinosauromorpha. These
are archosaurs that already looked almost like real dinosaurs, but still
differed in some features. Silesaurus had four long, slender legs and a
graceful build. It was a little over seven feet long and is now considered the
closest relative of the real dinosaurs. With over twenty of its skeletons
discovered, Silesaurus is the best-studied dinosauromorph. It has
contributed much to our better understanding of the early evolution of
dinosauromorphs.
Martin Qvarnström has also studied coprolites from this animal and made
exciting new discoveries. Silesaurus had long been thought to be an
herbivore, but Martin and his team discovered through its coprolites that
Silesaurus was actually an omnivore. In its fossilized feces, they found the
remains of many chitinous carapaces of beetles. The fact that we can make
such exciting discoveries in coprolites at all is thanks to computed
tomography and powerful synchrotron equipment. In the past, it was
necessary to cut open coprolites with a stone saw to see what they
contained. From the outside, it is usually impossible to see if and which
kind of food remains are hidden inside the coprolites. When cutting the
stones open, there is of course a risk of destroying their contents or cutting
the coprolite in the wrong place, so that nothing at all can be seen on the cut
surface. With microtomography, however, we can see all the details in
incredibly fine resolution without having to destroy the fossil.
These scans show us that Silesaurus hunted insects. Perhaps it did this by
stirring up leaves on the ground and then quickly snapping at them when a
beetle appeared. This strategy can still be seen in birds today. Although the
coprolites contained beetles, suggesting that Silesaurus mostly ate insects, it
is likely that it also consumed plants. This is indicated by its teeth, which
were distributed very irregularly and less numerous than in the jaws of most
other herbivores. But the animal is interesting in other ways, too, because it
tells us how the open ball-and-socket joint on the dinosaurs’ pelvis evolved,
and, in the future, could possibly reveal when archosaurs developed
feathers.
The oldest evidence of feathers in the fossil record is about 150 million
years old. These feathers belong to the Bavarian “Urvögel,” which I will
talk about in detail later. It is unclear at the moment if dinosauromorphs
already had feathers, because so far, we have no evidence for them. It
would be exciting to know at what time the feathers developed, because
even if there is no evidence for them so far, it does not necessarily mean
that feathers were absent in dinosauromorphs. Studies of the internal bone
structure of Silesaurus revealed that, just like the dinosaurs, it had a rapid
growth rate and a very active metabolism. Thus, it was able to maintain a
constant internal body temperature. Feathers would have helped Silesaurus
better retain this heat.
Silesaurus used to be considered a so-called facultative biped, because it
was believed to be able to choose between locomotion on all four or on two
legs. Most other dinosauromorphs were believed to be exclusively bipedal.
However, the latest research suggests that Silesaurus and its relatives were
quadrupeds throughout, because their forelimbs were significantly longer
than previously thought. This means that bipedalism, or movement on two
legs, did not evolve until the appearance of the real dinosaurs. In addition,
recent findings show that Silesaurus’s limbs looked quite different from
those of dinosaurs. Its thigh bones were straight, but they were connected to
the sides of its hips. This is different from dinosaurs and mammals. And
because the condyle of the femur in Silesaurus was neither inclined nor
fitted into an open socket, and the legs were not under the body, these
animals were not true dinosaurs and consequently were not ornithischian
(bird-hipped) dinosaurs, either. In fact, it is the open socket that
distinguishes dinosaurs from other animals. This evolutionary step must
have occurred only after the emergence of the silesaurs and shows that the
first true dinosaurs were more innovative than previously thought. Although
Silesaurus walked on four legs, its front legs were used more for support
and less for locomotion. It was only a small step before the first dinosaurs
balanced their bodies so that they could move bipedally. This offered the
advantage of being able to run faster than on all fours, and their heads were
higher off the ground this way. At the same time, this freed their hands and
opened them up for new uses; for example, they could reach for food or use
them as weapons. It also explains why the real dinosaurs eventually
outcompeted the dinosauromorphs, as their very special, agile mode of
locomotion is considered one of the key innovations that helped the group
to succeed.

Where the Wild Things Are

The rise of the dinosaurs was an important event in the history of

vertebrates. However, the exact time of emergence and the early split of
dinosaurs from other dinosauromorphs is not entirely clear. It is often
difficult to distinguish between dinosauromorphs and dinosaurs, as we have
already seen with the example of Silesaurus. But there is an animal from
Tanzania where the question of affiliation is even more difficult to answer.
It is called Nyasasaurus and was first scientifically described in 2013. The
genus name refers to Lake Malawi, which is also called Lake Nyasa.
Nyasasaurus was either the oldest dinosaur or the most closely related sister
taxon of this group.

In biology, a taxon is a name for a group of living organisms.

This group is assigned a specific rank, such as a species,
genus, family, or order. The International Code of Zoological
Nomenclature governs which scientific name is correct for a
particular taxon.

The animal has a unique combination of dinosaur-like characteristics and

already shows an increased growth rate. The study of Nyasasaurus indicates
that the global dispersal of dinosaurs was probably very slow at first.
Remains of the animal are from the Middle Triassic and are about 15
million years older than the second oldest dinosaur from that region. A 2013
phylogenetic analysis shows that Nyasasaurus is indeed a dinosaur.
However, although it is related, it is still outside all other dinosaur groups; it
is neither a true saurischian dinosaur nor a true ornithischian dinosaur. But
since the skeleton of Nyasasaurus is incomplete, further finds could still
change the classification, which is why the exact time of the origin of the
dinosaurs and the pace of their diversification remain unclear to this day.
Nevertheless, there is growing consensus among paleontologists that
dinosaurs were already primordial, albeit rare, components of Early Triassic
terrestrial ecosystems and that their ascent was slow and, in most cases,
regional. The oldest dinosaurs that can be reliably dated and identified are
from the early Upper Triassic of Argentina about 230 million years ago. By
the end of the Carnian, all three major dinosaur lineages—the Ornithischia,
the Sauropodomorpha, and the Theropoda—had evolved. Among the oldest
dinosaur finds worldwide are Eoraptor, Saturnalia, and Panphagia from
South America, which have been assigned to the sauropodomorph lineage.
The report on the oldest real dinosaur from Africa, Mbiresaurus, was only
published in 2022. Mbiresaurus comes from Zimbabwe.
Sauropodomorpha was the first major dinosaur group to spread and
diversify during the Triassic. During this time, many dinosaurs underwent
major changes in their body structure, becoming herbivores, increasing in
size, and switching to four-legged locomotion. By the end of the Late
Triassic, about 30 million years after the dinosaurs emerged,
sauropodomorphs dominated the niche of large herbivores in all continental
ecosystems worldwide. There were different lineages with widely varying
body sizes, from bipedal species weighing about ten kilograms to
quadrupeds weighing more than five tons, plus animals with different
feeding biomechanics. Currently, there are about fourteen different species
of sauropodomorphs from the Upper Triassic strata of northwestern
Argentina and southern Brazil. This is about half of all species known
worldwide for the Triassic. These finds have significantly improved our
knowledge of the sauropodomorph fauna over the past fifteen years. These
South American sauropodomorphs are important for understanding the
origin and early diversification of this group, showing us what helped the
dinosaurs achieve their preeminence within the Late Triassic group of
terrestrial vertebrates.

The First Giants:

Hundreds of Long-Necked Dinosaurs Stuck in Mud

The oldest known dinosaurs may have come from the Triassic in South
America or Africa, but there are remarkable finds in Europe as well. One of
the most interesting and probably largest European sites for dinosaur
remains is located in northern Switzerland, near the Rhine River, less than
six miles from the German border. Dating from the middle stage of the
Upper Triassic, sometime between 228 and 208.5 million years ago,
hundreds of plateosaurs died near what would become the city of Frick, and
were preserved in the mudstone. Plateosaurus was an herbivore that could
grow up to thirty-three feet long and weigh up to four metric tons, the
largest animal of its time in Europe.
Plateosaurus had a small head that sat on a long, flexible neck with ten
cervical vertebrae. Its body was reminiscent of the large long-necked
dinosaurs (sauropods) of which it was an ancestor. Its torso was stocky, and
it had, similar to its descendants, a long, flexible tail. It carried its long legs
vertically under its body. Plateosaurus could walk on all fours, but usually
moved bipedally. When doing so, only its toes touched the ground, but not
the soles of its feet, indicating that it could, at least at times, walk quickly
on two legs. The real long-necked dinosaurs were no longer able to do this.
The biggest difference to the sauropods were the short forelimbs, which
were about half as long as the hind legs. With its short arms and its hands
with long claws, it could grasp and hold large objects. Its chest and shoulder
girdle were relatively narrow.
When I visited the outcrop in the active clay quarry in Frick in 2011
during a field trip, I met a former fellow student from Mainz who was in the
process of recovering a new, nearly complete skeleton of a plateosaur.
Plateosaurus was probably the most common dinosaur in the Late Triassic
and there are more than fifty localities in Central Europe where it was
found. Since many complete skeletons of Plateosaurus have been found, it
is probably one of the best-known Triassic dinosaurs.
The first remains of a Plateosaurus were discovered in 1834 by the
chemistry professor Friedrich Engelhardt near Nuremberg. He passed the
find on to Hermann von Meyer, who is now considered the founder of
vertebrate paleontology in Germany. Von Meyer wrote in a letter to the
journal Neues Jahrbuch für Mineralogie, Geologie und Paläontologie on
April 4, 1837:
“Dr. Engelhardt in Nuremberg brought to the Assembly of Naturalists in
Stuttgart some bones of a giant animal from a breccia-like sandstone of the
upper Keuper of his region. This find is of great interest. The bones derive
from one of the most massive lizards, which is related to Iguanodon and
Megalosaurus because of the heaviness and the hollowness of its limb
bones. These remains belong to a new genus, which I call Plateosaurus; the
species is Plateosaurus engelhardti. The detail of this I will make known
later.”
Von Meyer recognized the similarity of Plateosaurus to the lizards of the
Mesozoic period reported from England. When he named the animal in
1837, the term “dinosaur” did not yet exist—the British paleontologist Sir
Richard Owen would only introduce it in 1841. Plateosaurus was thus the
first dinosaur described in Germany and one of the first ever to be given its
own name. At the time, only four other dinosaur genera were known
worldwide. When the first finds of Plateosaurus were made in the middle
of the nineteenth century in Trossingen in southern Germany, Friedrich
August von Quenstedt, another contemporary paleontologist, gave
Plateosaurus the nickname “Swabian lindworm,” in reference to the
dragons of Germanic mythology.

Although there are more than fifty sites in Germany, and many more in
France and Switzerland, most of the complete skeletons and much of the
exposed material come from only a few outcrops. The most important three
are Frick in northern Switzerland, Trossingen in southwestern Germany,
and Halberstadt in central Germany. The depositional conditions of these
sites are very similar, which is why the three sites are also described as
“Plateosaurus bone beds.”
These deposits extend over tens of thousands of square feet. In Frick, the
bone field is at least one mile long, and a new skeleton appears around
every thirty feet. It is interesting that there are a large number of complete
Plateosaurus skeletons at this site, but that all other vertebrates are missing
there. The only exception is a few primitive turtles of the genus
Proganochelys found here and there among the plateosaurs. The outcrops at
Frick, Trossingen, and Halberstadt are particularly intriguing because the
skeletons of the plateosaurs there are stuck in the sediment in an upright
position. However, the hind legs of many specimens are bent, and the thighs
are spread wide so that the knees point to the side. In this position, the legs
look a bit like those of frogs. Because of this posture, it was clear from the
beginning that the animals, after their deaths, were not further transported
on by rivers but buried by the sediment, on the spot. But how did these
plateosaurs die? Why are the legs bent in many specimens? And why do we
see the same posture in numerous animals at three different locations
hundreds of miles apart?

A Plateosaurus skeleton squatting in a frog-like posture in Frick, Switzerland.

PHOTO BY THE AUTHOR

The fact that plateosaurs are so abundant in the fossil record indicates
that they were the most common herbivores in their ecosystems. They were
often found in mass assemblages, indicating that they moved in herds and
roamed large areas. The layers where the Frick fossil deposit were found are
referred to as playa strata. The word playa comes from the Spanish word for
“beach.” In geology, however, playas are considered plains within mostly
drainage-free basins. These basins may be located in valleys between
mountains, for example, or they may be coastal plains. Over time, they are
gradually covered by layers of saline clay or marl. When it rained or the
water table rose, these plains would then turn into shallow salt lakes or salt
marshes. For the herds of plateosaurs, especially for the heavy specimens
among them that migrated across these playa plains, small, shallow gullies
in the dense mud of the marl formed veritable traps. The large animals got
stuck in the mire, while the juveniles could escape. The heavy plateosaurs
tried to pedal their way out of the mud, but their hind legs could not find
solid ground underneath the sticky mud, and, much like in quicksand, they
sank deeper and deeper until they finally gave up in exhaustion. The leg
muscles were then so weakened that the animals slumped down. The body
sank deeper into the mud and the powerless legs were spread apart. While
the animals were helplessly stuck in the mud, small predatory dinosaurs
came and started chewing on their upper bodies, which were sticking out of
the mud, regardless of whether the plateosaurs had already died or were still
alive. These predators were not big killers like Tyrannosaurus, but actually
tiny animals that under different circumstances would never have been able
to take down a plateosaur more than twenty-six feet long. Although we had
not been able to find any bones of these predators until recently, we knew
they were there, as evidenced by hundreds of shed or broken small
carnivore teeth in the Plateosaurus bone beds. The carnivores preyed on the
defenseless plateosaurs in their mud traps, so much so that the upright
animals embedded in sediment were often missing their heads, necks, and
shoulders.
However, there is a second form of deposition in which complete
skeletons in a recumbent position have been preserved. In these specimens,
the neck and tail are strongly bent over the back, indicating desiccation and
mummification after death. During heavy rains, these mummified carcasses
were then carried away by mudflows and redeposited in deeper places.
Over time, the mud dried out and settled. This and the load of new sediment
flattened the brittle bones of the plateosaurs. The dried soft tissues such as
skin, muscles, and tendons were not preserved and they gradually
disintegrated.
Although plant-eating plateosaurs and carnivore teeth have been found in
the Frick clay pit as early as the 1960s, it wasn’t until 2006 that an amateur
paleontologist finally discovered the first skeleton of a carnivore in a higher
stratum. Unfortunately, this skeleton was missing its head, but that was
found three years later during a closer examination of the surrounding
rocks. In 2019, the animal was described and given the scientific name
Notatesseraeraptor frickensis. The genus name Notatesseraeraptor means
“the predator with mosaic features” and refers to the fact that this carnivore
combines different features of different predatory dinosaur clades. The
species name frickensis refers to the place of discovery, the community of
Frick. Although dinosaur remains and their footprints have been found in
Switzerland for many decades, Notatesseraeraptor is the first carnivorous
dinosaur described from this country. Considering that there are hundreds of
Plateosaurus skeletons from Trossingen, Halberstadt, and Frick, but hardly
any other dinosaurs, the discovery of the Swiss Notatesseraeraptor seems
almost like winning the lottery.
Similar to the predatory dinosaurs, juveniles of Plateosaurus are virtually
absent from the playa deposits; they were not found until fifty years later at
Frick. The smallest specimens measured about sixteen feet. Perhaps smaller
animals had been light enough to escape the mud trap of the Plateosaurus
bone beds. Martin Sander, my former professor from Bonn, even claimed
that no young animals would ever be found in Frick. He had led a large-
scale excavation campaign there as a young man and contributed
significantly to deciphering the causes of the plateosaurs’ deaths. For more
than twenty years, his prediction seemed to be correct. Then, however,
something completely unexpected happened that both disproved and
confirmed his prophecy. For her master’s thesis, one of Martin’s students
described several vertebrae of a plateosaur from Frick. The vertebrae were
not yet completely fused and could therefore clearly be assigned to a
juvenile animal—however, they were huge, much larger than those of some
adult animals. Nevertheless, there was no doubt that they were the same
species. At first, no one could make sense of it. Then they took a closer look
at the internal bone structure of this animal and other plateosaurs. It turned
out that these dinosaurs apparently did not yet have the same active
metabolism as their descendants. They simply grew quickly when it was
warm and there was plenty to eat, and more slowly when there were not
enough resources. So, these animals showed a certain developmental
plasticity in their body growth. This means that you cannot necessarily tell
the age of the animal by its size. This young animal was not yet fully grown
but was already quite a bit larger than many of its adult conspecifics. And
although it was only a few years old, it was sinking into the mud because its
body weight exceeded the critical threshold. So, contrary to Martin’s
prophecy, there was a juvenile among the victims after all, but this animal
was more than sixteen feet long and certainly already weighed more than
one metric ton.
It remains to be seen whether Frick will surprise us again in the future
with new finds that will force us to rethink our current views. The chances
for that are good, because the bone bed seems to extend much farther than
previously thought. Only recently, more plateosaurs were discovered during
groundwork for a new residential area on the other side of the valley. With
the numerous finds of recent years, Frick is now considered one of the most
fossiliferous dinosaur sites in Europe. Halberstadt in Saxony-Anhalt will
not be able to compete with Frick for this title. Although there were
extensive excavations there in 1910 and 1914, the site was later built over
and is no longer accessible.
The first dinosaur bones in Trossingen, in Baden-Württemberg, were
found as early as 1904. The first excavations began there in 1911, and two
complete skeletons found there were then displayed in the Natural History
Museum in Stuttgart in 1913. In the 1920s, a second excavation took place,
in which German and American colleagues worked together. They
recovered a total of twelve skeletons, one of which was exhibited in the
American Museum of Natural History in New York. A third excavation took
place in the 1930s, but unfortunately, two thirds of the recovered finds were
destroyed during World War II.
In addition, there are dozens of smaller sites in Germany, many in
Franconia near Nuremberg, where over a thousand individual bones have
been found. Other famous sites are in Burgundy, France, and there are even
reports of a relative of Plateosaurus from Greenland. Even in the North
Sea, off the coast of Norway, isolated bones of Plateosaurus have been
discovered during an oil drilling, eighty-five hundred feet below the
seafloor. In the Upper Triassic, Plateosaurus was probably widespread
throughout Europe, and had closely related genera on every other continent,
with the exception of Australia. It is not surprising that these animals were
found worldwide, considering that in the Triassic, all continents were still
connected and formed the primeval supercontinent of Pangaea. Even in
Argentina, relatively large relatives of Plateosaurus, such as Riojasaurus
and Lessemsaurus, lived in the Upper Triassic. They were even a bit larger
than Plateosaurus and, with their length of up to thirty-nine feet,
foreshadowed the even larger sauropods that would succeed them in the
Jurassic. In the Triassic, however, these prosauropods were still the largest
animals in their respective ecosystems. At least that is what was believed
for a long time—until an extremely unusual discovery was made recently,
not in the field, but in the collection of the Tübingen Museum of Natural
History. There, the remains of a “plateosaur” discovered in 1922 were
reexamined more closely. It was found that in reality it was not a plateosaur,
but in fact a completely new species, much more closely related to the
sauropods than to Plateosaurus. The new name is Tuebingosaurus. This
finding could have far-reaching consequences, forcing paleontologists to
take another look at the “plateosaurs” in all the historical collections of
Germany, France, and Switzerland. Perhaps not all is Plateosaurus what
appears to be Plateosaurus.
At any rate, by the end of the Triassic, dinosaurs had evolved so much
that they became the true superheroes of their time.
And so the mass extinction that followed the Triassic offered the
dinosaurs infinite opportunities to evolve: a crisis as an opportunity.
The prosauropods evolved into giants and the carnivores became
increasingly dangerous predators. The legs they carried under their bodies
offered five advantages at once:

1. With two legs swinging vertically under the body, they could
run faster.
2. Being bipedal also allowed them to use their hands in other
ways. They could grab or hold on to things, and use their claws
as weapons to hurt or kill other animals.
3. The upright gait prevented them from dragging their bodies
across the ground, as is the case with lizards. This made the
dinosaurs less susceptible to overheating, even when the
ground was very warm.
 4. Their upright posture also enabled them to see farther into the
distance and detect potential danger or prey much earlier. This
gave them the opportunity to react preemptively, which proved
to be a great evolutionary advantage.
5. Their upright posture meant that they did not have to twist their
bodies sideways when walking, nor was the air forced from one
half of the lungs into the other. They could exhale it to make
room for fresh air. This improved breathing in an environment
that was less oxygen-rich than today’s.

However, elongated legs extending vertically under the body were not the
only key innovation of the dinosaurs. Their avian lungs, combined with
more efficient breathing, enabled them to walk with endurance and for long
distances. This allowed them to cover vast areas and disperse across the
world.
And sometime after splitting off from other dinosaur-like animals, such
as Silesaurus, the dinosaurs developed plumage. This was perhaps an
adaptation to a new catastrophe that struck the world. This time it didn’t get
hotter, but instead really cold, and the dinosaurs were prepared for it. The
end-Triassic mass extinction about 201 million years ago was one of the
five major extinction events in Earth’s history. In addition to the marine
conodonts, it caused the extinction of four major groups within the
crocodilian lineage that had previously been competitors of the dinosaurs:
the rauisuchians, the phytosaurs, the aetosaurs, and the ornithosuchians.
These large groups were similarly successful in the Triassic as the
dinosaurs. Yet the causes of mass extinctions during this period are not
entirely clear. Some of my colleagues cite sea level fluctuations. In
addition, climatic changes must have accompanied the extinctions.
Normally, climatic fluctuations are very slow processes, whereas this
extinction event may have been quite abrupt. In less than fifty thousand
years, all conodonts and many crocodile and amphibian species were
extinct. Other possible causes include volcanism or an asteroid impact, or a
combination of both.
However, because the sea level was extremely low—it was at a low point
worldwide at the Triassic–Jurassic boundary—we must assume that the
water was bound elsewhere. In 2022, a study found that there is evidence of
moraines at the end of the Triassic in northern China.
 A moraine is material that is moved along by glaciers then
deposited and piled up as debris elsewhere.

This makes a climate catastrophe seem more likely, and glaciation of the
polar ice caps may have led to significant cooling.
The dinosaurs’ feathers offered a significant advantage in this harsh
environment. This allowed them to access the rich, evergreen Arctic
vegetation even in freezing winter conditions, or to hunt for animals that
needed these plants for food. Recurrent, violent volcanic eruptions,
resulting in volcanic winters with massively reduced solar irradiation,
caused mass mortality on land to which all medium-sized to large non-
dinosaurs fell victim. These reptiles were not sufficiently insulated and
were decimated as a result. They were replaced by dinosaurs that were
insulated and protected against the cold. With their plush plumage, these
animals had adapted well to cold temperatures and were able to spread
rapidly through the Jurassic by adapting quickly. This ecological expansion
also led the dinosaurs into regions that had previously been dominated only
by large non-feathered reptiles.

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 The Jurassic

(201.3 to 145 million years before present)

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 Chapter 3

The Dinosaurs of the Morrison Formation

The Bone Wars

Everything we know about dinosaurs today is owed to the many
paleontologists around the world who are constantly discovering and
studying new fossils. What’s more, the way dinosaurs are found is often as
interesting as the animals themselves. In my view, the most exciting story
about dinosaur hunters is about the rivalry between two Americans, Othniel
Charles Marsh and Edward Drinker Cope. Among other things, they
searched for dinosaurs in the Morrison Formation, one of the most famous
rock sequences in North America in which fossils have been discovered,
and it was they who first scientifically described Stegosaurus,
Brontosaurus, Camarasaurus, and Diplodocus. In the nineteenth century, in
the so-called Wild West, the two paleontologists engaged in a tense dispute
over the discovery of new dinosaur species. There were even shootings and
sabotage. The hotheads accepted that some of their finds would be lost to
science forever. The feud was fought so bitterly that the country’s
newspapers reported on it in detail and referred to the dispute as the Bone
Wars.
The events of that time were so wild and adventurous that one might
think they had been devised by an author of Wild West novels. Some one
hundred and twenty years before the blockbuster Jurassic Park drew
crowds to the movie theaters, this dinosaur spectacle occupied the media,
and people flocked to newsstands in droves to learn of the adventurous
developments of the Bone Wars. It was not a Hollywood story that thrilled
young and old, but the then-new realization that dinosaurs must have
existed. The discoveries of the gigantic bones of the prehistoric lizards
appeared on the front pages of newspapers all over the world, and in the
United States especially, the interest was enormous. For the first time,
reports were published of gigantic, bizarre monsters and their fossils, found
in Colorado, Wyoming, and Montana. These areas were almost uncharted in
the 1870s, and the army was fighting Indigenous peoples there, pushing
them from their ancestral homelands. The stories of dinosaurs, cowboys,
Native American tribes, and two eccentric professors were eagerly absorbed
by the readers. For not only the finds provided interesting material for
stories, but also the people who searched for the primeval lizards. The fact
that dinosaurs existed was still new information at the time. The first
dinosaur was described by geologist and paleontologist William Buckland
in England in 1824, and Sir Richard Owen, the famous British physician
and zoologist, coined the term “dinosauria” in 1842, by which time both
Cope and Marsh had already been born. The first American dinosaur was
found by paleontologist Joseph Leidy in 1858, and the fact that Othniel
Charles Marsh and Edward Drinker Cope launched a veritable hunt for
fossils a little later, with the hunters armed not only with shovels and chisels
but also revolvers and rifles, can be largely attributed to their extraordinary
personalities. Each of them wanted to be the first to discover the next new
species of dinosaur, and so colleagues became archenemies and discovery
sites were defended to the death. The two men ruthlessly covered up
outcrops with dirt, destroyed bone fragments, and in some cases blew up
entire sites just to hide their finds from other researchers.
Cope and Marsh first met in Berlin in the winter of 1863, where Marsh
was studying anatomy. Cope was on a trip across Europe, and both escaped
a draft and the horrors of the American Civil War by spending time abroad,
even though they were of military age. Even then, it helped to be rich.
While hundreds of thousands died on the battlefields back home, the two
gentlemen were able to study and travel in peace in Germany. Cope’s father
is said to have sent his offspring to Europe primarily because of an affair
that he thought was not befitting his son’s station and thus wanted to stop.
When Cope and Marsh first met, they were initially on good terms. They
were even said to have taken walks through Berlin together and later wrote
letters to each other, and they exchanged fossils, photographs, and
manuscripts.
However, they were very different. Cope was the eager autodidact from a
rich Quaker family with an astute academic mind, and the rather sedate but
extremely methodical Marsh came from a poor background, but had a rich
uncle who endowed him with enormous financial resources. This uncle was
George Peabody, a businessman, investment banker, and one of the great
philanthropists of his time. Thanks to his uncle, Marsh was able to attend
Yale University, where he studied geology and paleontology, a relatively
new subject at the time, and then anatomy in Germany. After the two met in
Berlin, it didn’t take long for tensions to grow and the first small intrigues
to develop between them. This was probably due in part to the fact that
Marsh had already earned two academic titles at the time but had published
only a few scientific articles. Cope, on the other hand, who was almost nine
years younger, could already boast thirty-seven scientific publications, even
though he had dropped out of school after the age of sixteen.
These two men contributed a lot to the research of dinosaurs and other
prehistoric animals. Cope was a brilliant mind, describing more than a
thousand new vertebrate species and publishing some fourteen hundred
scientific articles during his lifetime, a record that remains unbroken today.
Marsh, who was in his final year of graduate school when Charles Darwin
published his book on the origin of species, was one of the first scholars to
adopt Darwin’s theory of evolution in the United States. He was an early
proponent of the view that birds were closely related to dinosaurs and
theorized in 1877 that they were the descendants of these prehistoric
animals. The reason that he came to this conclusion is probably linked to
the fact that the prehistoric bird Archaeopteryx was first scientifically
described in Germany in 1861—just a few years before Marsh’s trip there—
and his money certainly made it possible for him to see the fossil for
himself. He was actually a geologist and not as well-versed as Cope when it
came to vertebrate anatomy, but he had good connections in influential
circles and was a shrewd strategist with political skills. As important as both
men were to dinosaur research, they were apparently difficult
contemporaries. Marsh was said to be consumed by ambition—
overreaching, unscrupulous, and egotistical. Cope was described as quick-
tempered and overbearing. Naturally, it didn’t take long for the first friction
to develop.
As soon as they returned to America, the two scientists were appointed to
different universities: Marsh got the first chair of paleontology in the United
States at Yale University, and Cope became a professor of zoology at
Haverford College, but only after his parents pulled some strings. He was
awarded an honorary master’s degree, without which he could not have
become a professor. Both scientists initially named a few new species after
each other, which is a great honor in paleontology. Cope named a fossil
amphibian Ptyonius marshii, and Marsh returned the favor by naming a
giant marine reptile Mosasaurus copeanus. But Marsh’s gesture was less
friendly than it appeared. For as it turned out, the mosasaur came from a
quarry in New Jersey where Cope had previously worked. He had shown
this quarry to Marsh in 1868, and Marsh, behind Cope’s back, had arranged
with the landowner that new fossils from there would be shown to him first.
Thus, the first description of fossil species quickly became a competition.
Cope considered Marsh intellectually inferior and claimed—initially
behind his back, later publicly—that Marsh would have never been able to
publish anything without his support. He also accused his colleague of
having used him only to take advantage of Cope’s intellect. It was all the
more humiliating for Cope when Marsh, of all people, made fun of him in
front of the whole world when, in reconstructing the marine reptile
Elasmosaurus, Cope had confused the tail and neck of the animal and
mistakenly placed the head at the tail end. Marsh used this gaffe to ridicule
the “prodigy” Cope.
Cope’s publication on Elasmosaurus appeared in 1868. After that, the
two disputants never spoke favorably of each other again, and from 1873 on
their arguments were downright hostile, accusing each other, among other
things, of incompetence. The real bone war then started in 1877, when
Marsh received a letter from a schoolteacher in Colorado. This teacher—an
amateur fossil collector—and an acquaintance had been hiking in the
mountains near the town of Morrison, where they were looking for
fossilized leaves in the sandstone. Instead, the two found large bones
embedded in the rocks. The teacher wrote in the letter to Marsh that the
bones were probably a vertebra and a humerus of a giant lizard. He sent
Marsh a total of about seventeen hundred pounds of fossilized bones. But at
one point, when Marsh did not respond to a letter right away, the teacher
also sent Cope some of the specimens. Marsh heard about this and
published his research findings as soon as possible thereafter. He also paid
the teacher a hundred dollars for the finds, which prompted the teacher to
write to Cope again, asking him to forward his specimens to Marsh. This
offended Cope immensely. However, in March 1877, Cope received bones
from a school principal in Cañon City, also in Colorado. These remains
were from a dinosaur that must have been even larger than the one Marsh
had described. It was a long-necked dinosaur that was given the name
Amphicoelias fragillimus. Amphicoelias is hardly known to us today,
because unfortunately Cope’s original material was destroyed or lost.
Among experts, however, his species is considered a hot contender for the
record for the largest dinosaur ever found. The animal has since been
renamed Maraapunisaurus, because Amphicoelias fragilimus was only
know from fragmentary material at the time, and while it was believed to be
a diplodocid sauropod, more recent phylogenetic analyses show that it was
in fact a primitive rebbachisaurid. Hence, the name had to be changed.
This incredible find of Marsh’s rival prompted Marsh to act swiftly.
When he learned of a huge bone bed northwest of Laramie in Como Bluff
from two workers from the Union Pacific Railroad Company, which was
building a transcontinental railroad across the States at the time, he sent one
of his men there to take care of the excavation. Como Bluff turned out to be
a gold mine for fossils; it was a veritable dinosaur graveyard. However,
shortly after, Cope learned of the site as well and assigned his own team to
search for bones there. In the process, each team sabotaged the other’s
progress as best they could. Marsh tried to use his money and political
connections against Cope and to prevent his publications. Cope enticed
some of Marsh’s men to work for him instead. Because both feared that
their workers might change camps and reveal discovery sites, they were
extremely discreet about the exact locations of their fossils. When Henry
Fairfield Osborn, then a student of paleontology at Princeton, visited Cope
to ask him where fossils could be found in the American West, the latter
politely refused to answer. Mr. Osborn would still go on to play a big role in
the discovery of another famous dinosaur, later, as we’ll explore later in this
book.
The most famous finds from the Bone Wars—and the most famous finds
from the Morrison Formation overall—all came from the Como Bluff site.
The excavation site is located in western Wyoming and contains a sequence
of Upper Jurassic sediments. It was from there that the following dinosaurs
were first described: Allosaurus, Stegosaurus, Apatosaurus, Brontosaurus,
and Diplodocus.
The work at Como Bluff also marked the climax of the disputes between
the two scientists, and the methods they used to try to outdo one other got
completely out of hand. The damage done to science with these reckless
actions is immeasurable. To defend himself against Marsh’s coercion of his
men, Cope even hired some gunslingers, and on one occasion the fossil
hunters of both camps threw rocks at each other. It was pure luck that no
one was hurt in the process. Even the legendary bison hunter William Cody,
who went down in history as Buffalo Bill, worked for Marsh at one time,
supplying him and his team with fresh bison meat.
The Bone Wars lasted over fifteen years, leading to a mess of names for
the extinct giants—and two broken men who died impoverished and still at
odds with each other near the end of the nineteenth century.

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 Chapter 4

Germany’s Dinosaurs

How, on an Island, a Giant Turned into a Dwarf

The dinosaur books I read during my childhood were always about the
same dinosaurs: Tyrannosaurus, Triceratops, Stegosaurus, Brontosaurus,
and Ankylosaurus. We know all these animals from discoveries in the
United States. Most of my books were by American authors and had been
translated into German, so unsurprisingly they described the faunas of the
Jurassic and the Cretaceous of North America. The spectacular finds from
China had not yet been made at that time, and reports about dinosaurs from
South America had not found their way into popular science books. As a
child, I had already heard about the discoveries of the first dinosaurs in
England more than one hundred and fifty years ago, and about the
expeditions to Africa that took place around 1910. But one learned nothing
about current research from these countries in German children’s books,
and very little about discoveries in Germany. The only exceptions were
Plateosaurus from the Triassic, and Compsognathus and Archaeopteryx
from the Jurassic. Because Germany was largely covered by the sea during
the Mesozoic, and dinosaurs lived exclusively on land, we mainly find
ichthyosaurs, marine crocodiles, crinoids, and ammonites. Both
Compsognathus and Archaeopteryx come from lagoon deposits. They did
not live there but were washed into these shallow marine areas. In such
sediments we normally do not find any larger dinosaurs, and instead find
fish, horseshoe crabs, and occasionally a pterosaur that tragically fell into
the lagoon. However, because the Jurassic Sea was relatively shallow in
large parts, there were islands in it, and Europasaurus comes from one of
these islands.
 A mounted skeleton of Europasaurus at the Mega 2015 exhibition in Makuhari Messe, Japan.
PHOTO BY THE AUTHOR

Europasaurus is the dinosaur to which I have the strongest personal

connection, and I’ve researched it more than any other. For my diploma
thesis, I examined the occiput and the inner ear of some long-necked
dinosaurs with the help of computer tomography, one of which was
Europasaurus. The original material of this animal is housed in the
Dinosaur Park Münchehagen, where I later worked for a while. I lobbied on
behalf of the park and sold a replica of a complete skeleton of
Europasaurus to Japan, which became part of a special exhibition east of
Tokyo. I got to travel to the opening, help set up the skeleton, give a talk,
and was featured in a short welcome video. The show was called Mega
2015, and the booklet accompanying the exhibition included a short article I
wrote about Europasaurus and the area where it was found.
 I have given more talks about Europasaurus than any other dinosaur, and
to the most diverse audiences. I spoke not only to exhibition visitors in
Japan, but also to a large number of international colleagues. I even gave a
talk as part of the Children’s University in Göttingen. That was particularly
fun for me. The event was called “Europasaurus: A dwarf among giants!—
Dinosaur researchers as detectives.” The children were very engaged with
the topic. They asked exciting questions about dinosaurs and fossils, and I
showed them some original bones of Europasaurus, all of which were very
small, although long-necked dinosaurs usually grew huge. Together we
considered whether they might be the bones of a juvenile animal and
concluded that they must instead be from an unusually small adult animal.
The explanation truly sounds like a detective story. Europasaurus holgeri
was first discovered in northern Germany. The genus name means “the
lizard from Europe,” and the species name honors the finder of the first
remains, Holger Lüdtke, a fossil collector from the region. The fossil was
discovered at the Langenberg quarry in the Harz Mountains, where
sediments from the Upper Jurassic, about 154 million years old, are
exposed. This quarry has turned out to be an exceptional fossil deposit. Its
limestone and marl layers were tilted by about sixty to seventy degrees as a
result of tectonic movements and now form a steep face. They have yielded
an abundance of unique vertebrate fossils representing a specialized island
fauna, from the Upper Jurassic, never described from anywhere else in the
world. I have been to this quarry a few times to look for fossils there.
Unfortunately, I did not find any dinosaur remains—but I did find a very
large turtle. Its carapace was at least thirty inches in diameter, and I could
not carry the heavy block in which the fossil was enclosed, not even with
two colleagues. The boulder lay on a huge pile of other boulders that had
been scattered across the hillside after a blast. Gravity helped us roll the
boulder down the slope, and at the base of the rock face, a quarry employee
loaded it onto a forklift and hauled it away. The turtle’s carapace, which
was clearly visible on the rock surface, had several bite marks. But it is
difficult to say who caused the circular wounds. There were many large
predatory marine reptiles in the Upper Jurassic, but the culprit was most
likely a crocodile, as remains of at least four different crocodile species
were discovered at the Langenberg quarry. In addition to crocodiles, turtles,
and dinosaurs, small mammal teeth, flying reptiles and lizards have also
been discovered there. However, the most remarkable find from the
Langenberg quarry is actually the Europasaurus. While some genera of
long-necked dinosaurs are among the largest land animals of all time,
Europasaurus represents the first unequivocal dwarf sauropod and it is only
known from this site. Histological studies by Martin Sander and his team
have shown that some of those specimens were already fully grown despite
their small body size. While close relatives of Europasaurus such as
Camarasaurus and Brachiosaurus reached lengths of more than sixty-five
feet, the maximum body size of Europasaurus was probably less than
twenty-six feet, with a body weight of about eighteen hundred to twenty-six
hundred pounds. In 2006, Martin Sander and his team concluded that
Europasaurus was an example of a so-called island dwarf. During the
animal’s lifetime, a sauropod population was apparently cut off from the
mainland and evolved into a dwarfed form in just a few generations under
isolation on an island in the Lower Saxon Basin, while its closest relatives
on the mainland remained giants.
 Island dwarfing is an evolutionary biological phenomenon in
which the size of animals living in an isolated ecosystem
decreases within a few generations. Mostly this happens on
islands, where these animals do not have to fear predation
pressure. However, this phenomenon can also occur in other
habitats such as caves, isolated valleys, or inaccessible
mountainous regions. Other examples of island dwarfing
include Homo floresiensis, a small-bodied relative of modern
humans that lived on the Indonesian island of Flores until
about sixty thousand years ago; the dwarf mammoth from the
Russian Wrangel Island off the coast of Alaska, where such
mammoths still existed until about thirty-seven hundred years
ago; and the dwarf goat Myotragus from Mallorca, which
became extinct about forty-five hundred years ago. Dwarf
hippos have also been extinct in Madagascar for about two
thousand years, although reports of individual sightings
support the assumption that this species survived somewhat
longer in remote regions of the island. An example of an
extant dwarfed species today is the Sumatran rhinoceros of
Indonesia. Island dwarfing can be caused by an evolutionary
advance in sexual maturity or by a complete cessation of an
animal’s individual development. Both can result in a
retention of juvenile characteristics throughout later life
stages, i.e., paedomorphosis.

The high number of individuals at different growth stages at Langenberg

provides a unique opportunity to study ontogenetic stages (developmental
stages) and the interspecific variability of this small sauropod. Such an
ontogenetic range cannot be found anywhere else in the world. The
excellent preservation of a large number of skull elements from at least
fifteen individuals and postcranial skeletal material from more than twenty
animals has allowed scientists to conduct a detailed phylogenetic study of
these Brachiosaurus relatives. It is an important prerequisite for
understanding the evolutionary history of this sauropod group, because as
the continents were in motion and drifting apart in the Upper Jurassic,
dinosaur faunas evolved differently in the south and north. In the process,
some European dinosaurs of this time were linked with contemporary North
American faunas, while others appeared to be associated with Chinese
faunas. Brachiosaurids, which include Europasaurus, were found on both
the northern and southern continents.
From the abundance of skull bones, we chose the best-preserved brain
case of Europasaurus and scanned it at the University of Bonn using a
computer tomograph and then created a 3-D model of the brain, cranial
nerves, and inner ear with modeling software. Since I had already had the
opportunity to perform such scans myself during my time at Bonn, I also
created these computer models.
In 2015, two colleagues and I presented our results at a meeting in
Dallas, Texas. For our research, the dinosaur park had loaned us a few
specimens in advance and neatly packed them in boxes that were liberally
lined with foam. The fossils had been stored on the premises of the
University of Bonn for quite a while, and when I later flew to Japan for the
special exhibition, I had to deal with this very fragile and delicate material
again. Some of the original bones of Europasaurus were to be part of the
exhibition. Fortunately, the valuable specimens were never harmed, because
I always handled them with extreme care. Until one day, unfortunately, a
mishap happened: I was in a hurry and ran with a suitcase from my office to
the computer tomograph, which was located in another building. As I
rushed down the stairs, I missed the last step and fell—and the suitcase
slipped out of my hand. It sailed through the air before my eyes while sheer
horror gripped me. I had banged my knee pretty badly, but at that moment I
felt no pain. The case contained the best-preserved brain case of a
Europasaurus, and when I opened it, I saw that a piece had broken off the
side. I was so startled that tears welled up in my eyes. This was a unique,
especially valuable piece, and I had damaged it. I was distraught and
immediately ran to Olaf, our preparator. He noticed that I was completely
upset and calmed me down. Since the piece had detached with a clean
break, Olaf had no trouble reattaching it with a special fossil glue.
 The results ultimately showed that the examination of the skull had been
worthwhile. Since its anterior part was incomplete, the olfactory bulbs and
parts of the cerebrum could not be reconstructed, but we were able to
accurately re-create the posterior region of the brain and most of the cranial
nerves. In addition, we were able to represent various vascular structures,
such as the lagena and inner ears, the animal’s organs of balance.
At the Langenberg quarry, we found remains of Europasaurus only up to
a certain stratigraphic horizon, and after that there were no more finds of it
at all. For a long time, it was unclear why Europasaurus suddenly
disappeared from the fossil record. In recent years, however, we seem to
have solved the mystery. In younger strata, only a few feet above the strata
where Europasaurus was found, large footprints appear in the fossil record
of the Langenberg quarry. They are probably directly related to the
extinction of Europasaurus and other island dwarfs found in the region. The
layer in which these footprints commonly appear is about sixteen feet above
the Europasaurus horizon. Stratigraphers who have studied this rock
sequence estimate that this corresponds to about a thirty-five-thousand-year
time difference. The current working hypothesis is that a temporary sea
level depression occurred, creating a land bridge between Sauropod Island
and the mainland. This allowed predatory dinosaurs, which must have been
very large, to invade the island. It is not quite clear which animals they
were, but the shape of the footprints and sporadic finds of claws indicate
that they must have been close relatives of Allosaurus, which we know
from the Morrison Formation. One possible candidate is the Neovenator.
Judging from the footprints, the animals must have been twenty-three to
twenty-six feet long. The invasion of these predatory dinosaurs caused a
faunal upheaval on the island. The carnivores that previously lived on the
island were rather small, and apparently there had been a natural balance
between theropods and europasaurs. For the giant Neovenator, however, the
miniaturized long-necks were easy prey. Europasaurs could not oppose it
and died out within a few generations.

The First Feather That Changed Everything: Archaeopteryx

In science, coincidences often play a central role, and one such coincidence
was the discovery of the prehistoric bird Archaeopteryx, first scientifically
described by Hermann von Meyer in September 1861. Von Meyer’s paper
was about a single feather of a prehistoric animal—only mentioning a
complete skeleton in a sidenote—that came from the same strata and had
characteristics of birds and reptiles. Literally, he wrote: “From our living
birds it shows some deviation.”
The text is remarkable because von Meyer wrote it less than two years
after Darwin’s book On the Origin of Species was published. In this book,
Darwin cited numerous proofs for his theory that all animal and plant
species must have descended from a common ancestor. While von Meyer
did not elaborate on Darwin’s book, he seemed to have already known and
accepted the theory. Darwin called the underlying process for the gradual
change in all animal and plant species evolution. He saw its driving force as
competition among species for resources and habitats, with natural selection
favoring those forms that are stronger or better adapted to their habitat.
With the publication of this book, a broad public was confronted for the
first time with thoughts on the origin of species that contradicted the
biblical story of creation.
What made Darwin’s theory so difficult to grasp at the time was the lack
of connecting links in the fossil record. Opponents of his theory argued that
one would have to find substantially more transitional forms from one to
the other species, should the species have actually emerged from each other.
The fact that now, with the discovery of the “Urfeder” (German for
“prehistoric feather”), there was actual evidence for Darwin’s theory of
evolution, immediately after the publication of his book, was certainly
helpful in making his case.
The feather and the first complete skeleton of Archaeopteryx from 1861
were followed by another spectacular find of the prehistoric bird in the
1870s. This fossil is on display at the Natural History Museum in Berlin and
considered one of the finest specimens. The first Archaeopteryx skeleton is
now in the Natural History Museum in London. A total of twelve prehistoric
birds and one feather have been found so far. In the case of one of these
animals, it is now clear that it is not an Archaeopteryx, but a small,
feathered predatory dinosaur that was probably more closely related to the
sickle-clawed raptors than to birds. The other specimens also show
differences that justify subdivision into different species. That is why some
of my colleagues now avoid calling these animals Archaeopteryx; they
simply call them “Urvögel” (proto-birds).
 The London specimen of Archaeopteryx from the Solnhofen Limestone on display at the Natural
History Museum in London, United Kingdom.
PHOTO BY THE AUTHOR

In 1860, British biologist and comparative anatomist Thomas Henry

Huxley proposed that birds were descendants of dinosaurs. In doing so, he
pointed to skeletal similarities between dinosaurs, Archaeopteryx—which
he called the “first bird”—and modern birds. Marsh came to the same
conclusion in the United States, and his initial descriptions of early
Mesozoic tooth-bearing birds such as Ichthyornis and Hesperornis
supported this thesis.
This view persisted until the end of the nineteenth century, when Danish
paleontologist Gerhard Heilmann rejected the theory in 1926 and wrote in
his influential book The Origin of Birds that birds could not be dinosaurs
because dinosaurs did not have a wishbone. In birds, the left and right
clavicles are fused to form the V-shaped wishbone. After that, there was
silence on the subject for almost forty years, and it was generally assumed
that birds must have had crocodile-like ancestors, until American
paleontologist John Ostrom discovered a small carnivorous dinosaur in
1964 that he named Deinonychus, which means “the terror claw.” We know
these dinosaurs as the sickle-clawed raptors from the Jurassic Park movies.
Ostrom noticed the unmistakable similarity of the skeleton to that of
modern birds. He then also examined the skeleton of Archaeopteryx in
detail, which led him to claim that Huxley had been right and that birds had
indeed evolved from dinosaurs. To honor Ostrom’s contributions, a
feathered predatory dinosaur from the Altmühl Valley was named Ostromia
in 2017. Ostromia is closely related to Deinonychus (a dinosaur which
Ostrom named). Ostrom concluded that birds were dinosaurs because
they’re so similar to extinct dinosaurs. Funnily enough, Ostromia had long
been mistaken for a “bird” (Archaeopteryx) which underscores how similar
birds and maniraptorans (the group to which Ostromia and Deinonychus
belong) really are. Incidentally, this Ostromia is the supposed
Archaeopteryx specimen that was actually a raptor—and its namesake is
therefore extremely apt.
Ostrom’s work and the subsequent publications of paleontologist Robert
Bakker initiated a veritable renaissance in dinosaur research and led to a
profound rethinking of almost all aspects of dinosaur biology. They created
the foundation for our current image of dinosaurs and for the way these
animals are portrayed in film and on television. And it all started with the
discovery of a single Archaeopteryx feather.
However, the discovery of Archaeopteryx is not only historically
important. Especially the last three specimens discovered have contributed
to our better understanding of these animals today. Oliver Rauhut and his
team from Munich University were also able to distinguish them from
another small predatory dinosaur that was also discovered in the Plattenkalk
(a very finely grained limestone). They dubbed it Alcmonavis, “the bird
from the Altmühl River.”
In the tenth specimen of Archaeopteryx, which is excellently preserved,
structures on the skull that could not be observed before were recognizable
for the first time, and the bones in its foot showed unusual features. In the
eleventh skeleton, the feathers are particularly well preserved, and it is
possible to clearly see the pennaceous feathers. The research team was able
to show that the feathers of Archaeopteryx were already asymmetrically
shaped, as in modern birds. What this means and how important this
discovery is will be explored in a later chapter, when we take a closer look
at the evolution of birds.
The latest fossil to date was discovered in 2010 in a quarry near the
Köschinger Forest north of Ingolstadt. It comes from strata of the so-called
Mörnsheim Formation and, at around 153 million years old, is the oldest
find of the prehistoric bird to date. The fossil is about as old as the strata
from the dwarf island of Langenberg. This makes my ears prick up, because
it is quite conceivable that one day we might also discover remains of a
proto-bird from the Langenberg quarry. Martin Sander predicted this several
years ago, and I am curious to see if he will be proven right. Should such an
animal actually appear one day, I would call it Sanderopteryx predicta
—“the predicted Sander’s feather.”

The Adventures of Werner Janensch

I grew up in the very south of Germany on beautiful Lake Constance and

only came to Berlin for the first time after the reunification of East-and
West-Germany. At that time, the first thing I wanted to do, of course, was to
visit the Museum für Naturkunde in East Berlin, especially to see the Berlin
Archaeopteryx specimen and the Brachiosaurus skeleton. Brachiosaurus, or
“arm lizard,” was a large long-necked dinosaur whose arms were longer
than its hind legs, quite unlike many other sauropods such as Diplodocus,
Brontosaurus, and Apatosaurus. The skeleton in the museum’s Great
Dinosaur Hall consists of original bones. To date, it is the largest complete,
mounted original skeleton of a dinosaur in the entire world. During its
lifetime, the animal must have been some seventy-five feet long and its
head towered about forty feet above the ground. Brachiosaurus was found
by the paleontologist Werner Janensch. Janensch was an eminent vertebrate
paleontologist, geologist, explorer, and one of the most important German
dinosaur specialists of his time. As a young man, he traveled to Africa to
search for dinosaurs. Strictly speaking, the skeleton—or perhaps I should
say the skeletons—of the Berlin exhibition is comprised of not just one, but
at least two dinosaurs of the same species. Because Janensch found several
partial skeletons that complemented each other well, it was decided during
the mounting of this reconstruction, to build a composite from bones of
various animals in order to be able to show as complete a skeleton as
possible.
At the end of the nineteenth century, Reich Chancellor Otto von
Bismarck, following the example of England and France, finally agreed to
acquire colonies in Africa after long urging by the imperialist movement in
Germany. One of these colonies was called German East Africa. It existed
from 1885 to 1918 and included the present-day countries of Tanzania,
Burundi, as well as Rwanda and some areas of Mozambique. The colony
had a total area of 384,000 square miles, almost three times the size of the
Federal Republic today. German East Africa was the largest colony of the
German Empire, with almost 8 million people living there by its end. After
the end of World War I, in accordance with the provisions of the Treaty of
Versailles, the colony was divided between Belgium and Great Britain and
placed under the administration of the League of Nations.
The young Janensch found himself in German East Africa in April 1909.
Three years earlier, a mining engineer there had discovered a huge bone
sticking out of the ground on the slope of a hill. The locals called the hill
Tendaguru, or the steep mountain. The Upper Jurassic rock sequence
exposed there is called the Tendaguru Formation. It is considered one of the
most important outcrops of Jurassic vertebrate fauna in the southern
hemisphere. The Tendaguru mound is located near the port town of Lindi on
the southeast coast of the former colony. Today, the town is part of
Tanzania. The mining engineer who made the find informed his superior in
Hanover, and he forwarded the report to the Commission for the
Geographic Exploration of the Protected Areas. This department of the
Imperial Colonial Office dealt with the geographic assessment of the
German protectorates. Hans Meyer, a German Africa researcher, heard
about the find through the grapevine, who in turn informed the
paleontologist Professor Eberhard Fraas. Fraas was asked to examine the
find on-site, and traveled to Lindi by steamship, arriving in August 1907.
After a five-day walk, Eberhard Fraas reached the Tendaguru mound. It
was immediately clear to him that these must be dinosaur bones. Fraas also
observed that the Tendaguru Formation was unusually rich in fossils. The
mining engineer who made the discovery joined the professor and, with a
local team, helped him recover two large skeletons of long-necked
dinosaurs and ship them to Germany. These giants were later described as
the genera Tornieria and Janenschia.
After his return to the German Reich, Fraas tried to raise enough money
for an expedition to Africa. He found support from Professor Wilhelm von
Branca, who was at the time the director of the Geological-Paleontological
Institute and Museum of the Royal Friedrich Wilhelm University in Berlin.
For von Branca, it was a matter of German national pride to help such a
project succeed. He collected a large sum of money from his wealthy
friends and sent his best man, curator Werner Janensch, to Africa. Janensch
arrived in Dar es Salaam in April 1909 and led four field campaigns until
1912. However, because the region around Tendaguru was infested with
tsetse flies, which transmitted nagana disease to horses and other pack
animals, Janensch had to rely on the help of many local porters. In the first
season, he employed about one hundred and sixty local workers. With their
support, he opened about a hundred outcrops and shipped the recovered
fossils to Germany.

In addition to the sauropods Tornieria and Janenschia discovered by

Fraas, there were other long-necked dinosaurs in the Tendaguru Formation.
One was the medium-sized Dicraeosaurus, which was relatively common
in these strata and is now also on display in the Berlin Natural History
Museum. But the find that made Janensch famous was a larger sauropod
that came to light around half a mile northeast of the hill. Janensch made
the discovery of the giant back in June 1909 and later named the animal
Brachiosaurus brancai. The first find consisted of a few tail vertebrae, the
sacrum, the pelvic bone, a few ribs, the upper and lower arms, shoulder
blades, femora, and the calf and shin bones. Although this was a juvenile
that was not quite fully grown, the humerus was already five feet three
inches long and the femur five feet one inch. A second find followed in
August 1909, when about one hundred and fifty bones of various dinosaurs
were discovered in a large outcrop, including two femora of Brachiosaurus.
In September of the same year, Janensch found more bones in a third pit,
widely scattered, along with bone fragments that were already heavily
weathered and could not be precisely identified. Finally, in October 1909,
he made his most important find, south of the hill, on the bank of the
Kitukituki River, where two skeletons of Brachiosaurus lay at once. At this
place the bones were so abundant that Janensch continued to dig there until
1912. He assumed that the animals must have sunk into the mud because
they were found in an upright posture with their arms and legs stuck
vertically in the sediment.
In total, Janensch must have found at least seven individuals. How many
there actually were is unclear. He did not keep accurate field notes and did
not document all the sites where they were found. In addition, some fossils
from Tanzania were destroyed by the Allies during an air raid on Berlin in
1943. Janensch had collected so much material that at that time not all the
boxes from Africa had been opened and examined yet, which is probably
why many a treasure was lost at that time. Janensch spent the rest of his
career examining and describing the finds. As he worked on the
Brachiosaurus material, he noticed that many elements bore a strong
resemblance to an animal from the Morrison Formation that American
paleontologist Elmer Riggs had described back in 1903. He had named it
Brachiosaurus altithorax. Janensch chose the same genus name for the
long-necked dinosaurs from the Tendaguru Formation, but added a new
species name, brancai, to honor Professor von Branca.
When paleontologist Gregory S. Paul examined the skeletons of
Brachiosaurus altithorax and Brachiosaurus brancai in 1988, he concluded
that it was justified to establish a subgenus for both animals. Paul was
aware of the historical significance of the names and suggested that the
animals be called Brachiosaurus (Brachiosaurus) altithorax and
Brachiosaurus (Giraffatitan) brancai. In practice, however, this designation
with subgeneric names was never used. The situation did not change until
Mike P. Taylor, another paleontologist, published an article in 2009
explaining that there were actually so many differences in these animals
that they needed to be given different generic names. He named Janensch’s
animal Giraffatitan brancai. I remember the article very well because I had
resumed my paleontology studies in Bonn shortly before it appeared. My
first lecture in the seminar was a summary of the article, and we discussed
whether this renaming was justified. Coincidentally, Mike Taylor was
visiting Bonn a little later, and I was also able to talk to him personally
about this issue. One of his arguments was that Janensch had probably
never seen the original bones of Brachiosaurus altithorax. However, I
wonder if this argument is relevant. I have seen both dinosaurs with my
own eyes, and they seemed very similar. In addition, Taylor argued that the
number of distinguishing features between the two animals was so great
that they should definitely be separated by name. I find this argument
understandable, though not tenable. Indeed, the International Code of
Zoological Nomenclature, which lay down the naming and classification of
all animal species internationally, do not specify the number of
characteristics in which one animal must differ from another in order to
belong to a different genus.
This is not trivial at all. To better explain, let me give an example. In
today’s animal kingdom there are the so-called true big cats, which carry
the scientific name Panthera. The big cats include the tiger (Panthera
tigris), the jaguar (Panthera onca), the lion (Panthera leo), the leopard
(Panthera pardus) and the snow leopard (Panthera uncia). All of these
animals look very different, live in different habitats, and some are of
different sizes and weights. Nevertheless, they belong to one and the same
genus. Nobody would think of splitting this genus. Just because the two
long-necks differ is not a sufficient reason to divide them into two different
genera.
Only if other dinosaur genera would stand between the two in the
dinosaur family tree would one need to change the name. But the only
dinosaur that might qualify for this scenario is Sonorasaurus thompsoni
from Arizona. In a 2016 study, Sonorasaurus is phylogenetically
sandwiched between “Giraffatitan” and Brachiosaurus, but in another
2017 study, “Giraffatitan” and Brachiosaurus are more closely related, and
Sonorasaurus is only a sister taxon to “Giraffatitan.”
The problem with the 2016 phylogenetic tree would be that the genus
name Brachiosaurus would be split and another genus would be stuck in
between. According to the rules of zoological nomenclature, this is not
possible. It would make the genus Brachiosaurus paraphyletic. This means
that Sonorasaurus would be split off from a larger group, even though it has
fewer differences, since it would lie between the other two, and the
remaining remnant, i.e., B. altithorax and B. brancai, would remain lumped
together in the absence of other visible features. This would indeed be
wrong and would not make sense. But even if we follow the 2016
phylogenetic tree, there are two possibilities: either we actually rename B.
brancai, or we would have to change Sonorasaurus to Brachiosaurus
thompsoni. From my point of view, the latter would be more justifiable,
since the name Sonorasaurus thompsoni has not been around that long. The
exact classification will perhaps become more interesting when the
brachiosaurid, found in Portugal in August 2022, is examined and
scientifically described. It is already considered the largest dinosaur ever
found in Europe. Which species it is, however, is still uncertain.

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The Lower Cretaceous

(145 to 100.5 million years

before present)

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OceanofPDF.com
 Chapter 5

Argentina—Where the Giants Live

The Long-Necked Dinosaurs That Tower over Everything

For many decades, the Berlin Brachiosaurus skeleton found by Werner
Janensch was the largest complete original skeleton of a long-necked
dinosaur in the world that could be admired in an exhibition. Other
candidates for the title of largest land creature mostly came from the United
States. But in the early 1990s, enormous bones of much larger giants were
found in Argentina. These giants belong to a group of long-necked
dinosaurs that are called titanosaurs, which survived into the Cretaceous
period. The very first finds of titanosaurs from South America were made
about one hundred and thirty years ago, but it was not until the 1990s that
remains of long-necked dinosaurs were found that stretched the boundaries
of what was deemed physiologically possible. With the description of
Argentinosaurus in 1993, the world was introduced to a dinosaur that must
have been almost 100 feet long and, depending on which author and which
calculation method one may follow, probably weighed between sixty and
ninety tons!
Sauropods are unique in many ways. We know of no comparable
creatures in the animal kingdom today. They have a build that does not exist
in any other group of animals. The obvious and most important feature is
their neck. No other animal has such a long neck as the long-necked
dinosaurs. Surely many will automatically think of a giraffe at this point
and suggest that giraffes also have long necks. However, this is relative—
the neck of an adult male giraffe can grow up to six feet long. This
corresponds to the average height of a human male. From our point of view,
that’s enormous. But some dinosaurs had necks up to forty-five feet long.
By comparison, the neck of a giraffe is downright tiny. To produce such an
extremely long neck, the sauropods developed first one and then a second
additional cervical vertebra and also shifted the shoulder girdle backwards,
so that functionally, the first dorsal vertebra became another cervical
vertebra. In addition, each cervical vertebra became significantly more
elongated, while the weight of the vertebrae was drastically reduced. This
was possible because the walls of the vertebrae were paper-thin and
interspersed with a complex network of air sacs. These air sacs provided
efficient respiration through unidirectional airflow, and also enabled the
sauropods to better remove excess heat and avoid overheating. This was
necessary because the long-necked dinosaurs, at least in the early stages of
their lives, had an active metabolism and grew rapidly. The rapid body
growth, the long neck, their avian lungs, and an efficient digestive tract
ultimately enabled the sauropods to grow so large.
The dwarfed Europasaurus is a special case among the long-necked
dinosaurs, because most sauropods were giants—true titans! No other land
animal was even close to that size. The African elephant, the largest land
creature today, is a dwarf compared to the “lizard-footed”—that’s what the
term “sauropod” means in English. The largest sauropods were ten times,
perhaps even twenty times, the weight of an elephant. One has to imagine
enormous colossi, some of which must have been as heavy as a jumbo jet or
a sequoia tree. These animals were mountains of flesh, like long-necked
blue whales on four legs. It is not without reason that the first sauropod ever
described was also named Cetiosaurus, “the whale-lizard.” I like this name
very much, because it is so pictorial. In my opinion, the long-necked
dinosaurs have the most fitting names.
One of the most beautiful names belongs to the first dinosaur I remember
learning about as a child: Brontosaurus, “the thunder lizard.” He is also a
long-necked dinosaur. As a little boy, I always imagined how the Earth must
have shaken with every step this giant took, and then I’d stomp through the
house with a loud roar, much to the chagrin of my parents and siblings.
Another giant was named Seismosaurus, “the earth lizard.” Later it turned
out that it was a particularly large Diplodocus species, hence the genus
name has since become invalid. Basically, most sauropod names contain an
allusion to their enormous size: Seismosaurus, Cetiosaurus, Brontosaurus,
Supersaurus, Ultrasauros, Notocolossus (“the colossus of the south”), and,
of course, Titanosaurus, namesake of the titanosaur clade. They get their
name from the Titans of Greek mythology. These were giants in human
form, a powerful race of gods. That paleontologists chose such names to
describe the enormous size of sauropods is understandable, especially when
you compare these creatures to extant animals. The smallest dinosaur of our
time can illustrate this impressively: bee hummingbirds (Mellisuga
helenae). These very beautiful birds live in Cuba, with the plumage of
males and females iridescent in different colors. The females weigh no
more than a penny (0.08 ounces) and the males less than 0.07 ounces,
which is about the weight of a gummy bear. Nevertheless, they are true
theropods; that is, they belong to the group of dinosaurs that also gave rise
to Tyrannosaurus, the king of all carnivores.
The largest sauropods are now known to have weighed at least fifty tons,
and many of my colleagues even consider body masses of over one hundred
tons possible. This means that the heaviest dinosaurs weighed 50 million
times more than the lightest members of this group! Fifty million times
more! That is fifty thousand multiplied by one thousand. But how does one
calculate the weight of a dinosaur? One of the methods for determining the
body mass of four-legged dinosaurs involves the circumference of the thigh
bone and the humerus. This is certainly easy to understand, because the
extremities have to carry and move the weight of the animal around. The
heavier the animal, the stronger the arm and leg bones must be to hold the
corresponding muscle mass. Most researchers agree that the crown for the
heaviest dinosaur currently still belongs to Argentinosaurus.
Argentinosaurus, however, is known only by its thigh, a few vertebrae and a
few small bone fragments. This femur is indeed massive, but I find it
difficult to accept it alone as a reliable reference for calculating its mass. In
general, most skeletons of the long-necked dinosaurs we know today are
extremely incomplete. In 2014, however, a team of researchers discovered
an entire herd of titanosaurs in Chubut Province, Argentina. I have a very
special relationship with these giants because I know the paleontologist
José Carballido, who recovered the animals and later scientifically
described them, very well. He worked as a postdoctoral researcher in Bonn
while I was studying for my degree. In 2014, when José discovered this
new colossus, I had just started a job as a scientific advisor for Dinosaur
Park Münchehagen. At the time, stern TV was reporting on the giant find,
and I had been invited to appear on the show on behalf of the park. While
we were talking about the very largest dinosaur ever found, I prepared a
fossiliferous block live on air that contained bones from the shoulder girdle
of the dwarf sauropod Europasaurus, the smallest representative of this
group. Since the dinosaur park had a subsidiary that produced life-size
dinosaur models, I was asked at that time to establish contact with the
museum in Argentina, and after long negotiations, the world’s largest live
reconstruction of a dinosaur was finally built under my guidance and
supervision. The model is truly gigantic. Originally, it was supposed to have
been exhibited in an old warehouse in Trelew, Argentina. But since the
animal was too big for the hall, the model now stands in the open, outside
the city near Trelew International Airport.

Many more bones are preserved from this titanosaur than from
Argentinosaurus, including the humerus and femur. To reconstruct the
anatomy of the animal, bones from at least three individuals of similar size
were examined. José then gave the dinosaur the scientific name
Patagotitan, “the Titan from Patagonia.” After applying various methods, a
reliable mass estimate of about sixty-nine tons was obtained. However, the
standard error in such a calculation is about seventeen tons. So the animal
could actually have weighed between fifty-two and eighty-six tons! This
sounds like a large margin of error, but it is relatively small compared to
estimates for other sauropods. We have a great many skeletal elements of
Patagotitan available and can use several different measurement methods,
all of which confirm a similar weight for the titanosaur. For
Argentinosaurus, such mass estimates cannot be calculated with scaling
equations because of the lack of the humeri, which makes the standard error
much larger for it. In addition, the anterior dorsal vertebrae of
Argentinosaurus are about 10 percent smaller than those of Patagotitan.
Thus, Patagotitan actually represents the largest known dinosaur species.
Probably more finds of giants are to be expected from Chubut, and I would
not be surprised if one day an animal is discovered there with a size that
would put even Argentinosaurus and Patagotitan in the shade.
The museum in Trelew has made many casts of the bones of Patagotitan,
and replicas of this giant are now located all over the world. In the Field
Museum in Chicago, there is a model in the main hall that is about one
hundred and twenty feet long and over twenty-eight feet high. In late
February 2022, during a trip to Argentina, I visited the workshop in Trelew
that makes these giants. At the time, I was to participate in a campaign by
Professor Oliver Rauhut from Munich, who researches and was searching
for dinosaurs from southern Argentina. This was at a time when
international air travel was extremely restricted due to the Covid pandemic.
Oliver didn’t know if this campaign would even take place and couldn’t
confirm it until a week before my flight. Fortunately, my Covid test came
back negative, and a short time later I was already on the plane from
Frankfurt to Buenos Aires. After a one-day stopover, I continued to Trelew
the next morning, where Oliver kindly picked me up from the airport.
Even at the Trelew airport, you feel as if you’ve landed in dinosaur
country. On the wall of the baggage claim area, a relief of a dinosaur
excavation site with bones of the prehistoric giants is emblazoned, and a
little farther on there are posters of dinosaurs, advertising the local
paleontological museum. It is a major attraction of this city and brings in
many international visitors. Its exhibition is worth seeing, but the real
treasures are in the collection, which is not open to the public. Among other
things, the original bones of Patagotitan are kept there. One night I stayed
at the Touring Club Hotel. The name sounds like it’s a flophouse for
backpackers, but it’s actually a legitimate hotel with over one hundred and
twenty years of history. Supposedly, the famous outlaws Butch Cassidy and
the Sundance Kid stayed here for some time after fleeing the United States.
The next morning, we loaded up the two SUVs and headed west. We
drove about four hours on National Route 25 to the village of Paso de
Indios. Along the way, we saw many impressive Cretaceous and Jurassic
rock formations, and the landscape often reminded me of the American
Midwest. In Paso de Indios, we refueled and bought some provisions before
continuing north. Route 12, one of the main north-south connections in
Chubut, is an unpaved dirt road, and I soon realized why Oliver had chosen
the off-road vehicles. His goal for this campaign was to investigate an older
outcrop where parts of a long-necked dinosaur skeleton had come to light
some time ago. He wanted to find out how much of the skeleton was still
hidden in the rock so that he could then decide whether to recover it later
with heavy machinery. Another focus of this campaign was to find new
outcrops. Oliver has been working in this part of the world for over twenty
years, but it was my first trip to Argentina and South America. I was
completely overwhelmed by the beauty of its environment and the many
animals and plants that were foreign to me. Most impressive to me were
certain large insects, grasshopper relatives, which do not exist here in
Europe, and the many birds of prey with their remarkable wingspans. But at
the sight of the nandus and guanacos I fully realized that I was in an
absolutely strange world. Except for in Kenya, I had never seen such large
flightless birds in the wild as those, simply running through the wilderness
here. These nandus sprint seemingly effortlessly, at up to thirty-seven miles
per hour across the rocky terrain. The guanacos are a wild species of the
llama genus, though the llama we know is a domesticated species. You just
don’t get to see animals like this in the wild in Europe.
We continued on Route 12 north along the Rio Chubut until we reached a
small village called Cerro Cóndor. There were only a few individual houses,
as well as a school that was apparently attended by the children of all the
surrounding farms. Without the school, the village would probably no
longer exist. Next to it was a small museum with some dinosaur finds from
the surrounding area. Directly in front of the village was an outcrop of
pterosaurs, which one of Oliver’s doctoral students wanted to study more
closely. A few miles farther north, about half an hour’s drive away, we
stopped to look for new dinosaur sites. The rocks there are assigned to the
Cañadón Calcáreo Formation and are between 160 and 150 million years
old. Thus, they range from the Oxfordian to the Tithonian, the oldest and
youngest stages of the Upper Jurassic, respectively. Even if the giant
titanosaurs of Argentina do not come from these strata, the rock is
immensely exciting for us. Unlike the Upper Jurassic in North America,
Europe, and Asia, we know little about this time in the southern
hemisphere. Since the contest between Cope and Marsh, many North
American Upper Jurassic sites have become known and been well studied,
but from the southern hemisphere we only know the sites in Tanzania.
Oliver has made it his mission to better study this period of Earth’s history,
and this is important because we can already see that long-necked dinosaurs
evolved quite differently in the northern hemisphere than in Gondwana, the
southern hemisphere continent. Furthermore, in no other country on Earth
can we trace sauropod evolution in the fossil record for as long as in
Argentina. Some colleagues even suggest that the cradle of sauropods was
in Argentina. In fact, the oldest long-necked dinosaurs are already found in
strata from the Norian of Argentina, a stage of the Upper Triassic that is
about 228 to 208.5 million years old. Sauropods are still missing completely
in the fossil record of North America and Europe from this time. But
already in the Jurassic, these giants existed everywhere on Earth, and while
their history was largely uniform into the Upper Jurassic, evolution in the
Cretaceous progressed completely differently in the northern and southern
hemispheres. In the United States and Canada, where long-necked
dinosaurs were long the dominant group of herbivores, they became rare at
the beginning of the Cretaceous until they then disappeared completely
from Colorado, Wyoming, Montana, and North and South Dakota. Only in
the far south, in the region around southern Texas and New Mexico, small
isolated colonies of long-necked dinosaurs remained in some cases. This
was a species of titanosaur called Alamosaurus, while the long-necked
dinosaurs in the north were successively replaced by the duckbill dinosaurs
and the horned dinosaurs.
The situation was quite different in South America, where these
ornithischian dinosaurs were almost absent. But even here, the Diplodocus
and Brachiosaurus relatives were gradually replaced by titanosaurs, which
attained an amazing species diversity. There were tiny titanosaurs—which
sounds like an oxymoron—medium-sized forms, and huge giants; there
were long-necked and short-necked forms, grazing animals and those that
ate foliage from the tallest trees. Sauropods occupied a wide variety of
ecological niches and thus prevented the development of other species. But
in the Jurassic Cañadón Calcáreo Formation, north of Cerro Cóndor,
titanosaurs were not yet present, and we therefore hoped to find other
earlier forms of long-necked dinosaurs there.
 On our quest for Jurassic dinosaurs, we drove through stunning landscapes in Chubut Province,
Argentina.
PHOTO BY THE AUTHOR

It’s always a pleasure for me to be part of a field trip and to excavate

fossils that have lain dormant in the sediment for millions of years—but it’s
even more impressive when you discover your own outcrop with dinosaur
remains. Sure, I helped dig at the pterosaur site, and I even helped dig up
some long-necked dinosaur bones; but those outcrops had been discovered
by others. I wanted to make my own discovery, to find a treasure myself
and be the first to excavate fossil bone fragments in the middle of nowhere
that could potentially deepen our understanding of the lives of those
prehistoric giants. However, the search for these remains is not always
crowned with success, even in an area as rich in fossils as Chubut Province.
The area is simply very vast and you could sometimes wander around for
days without finding anything. This can be frustrating, and that’s what
happened to me on this trip—that is, at least, until the last day! We had
driven north in our SUVs and parked the vehicles near the river. Around ten
o’clock, each of us headed off in a different direction; we had arranged to
meet back at the car at 5:00 p.m. I hiked over hills and through small
canyons, climbing steep slopes, always keeping my eyes on the ground in
front of me. I turned over every stone that even remotely looked like a bone
or shimmered bluish, because in this area the fossilized bones often have a
pale blue coloring. Unfortunately, bluish lichens also grew on the stones
here, so I often reached out in vain. In the afternoon, after having found
nothing for hours, I finally stumbled upon a piece of eggshell from a nandu.
It was lying on the scree slope of a larger hill, and I wondered if it might
have come from an abandoned nest from up on the peak of the hill. Perhaps
I could find another larger eggshell there to bring back as a souvenir for my
children. I climbed up, but I couldn’t find a nest anywhere. Instead, out of
the corner of my eye, I saw a small bluish stone. Instinctively I picked it up,
although I assumed that it was just another stone covered with lichen. But
instead it was actually a piece of a fossilized bone. I wiped the dust off the
surface and did what any paleontologist does when he wants to make sure a
fragment is a bone in the field: I held the fragment’s fractured surface to the
tip of my tongue and licked it carefully. The interior of fossil bone consists
of very porous material, and if you touch this surface with its tiny pores
with your wet tongue, a slight vacuum is created and the bone sticks to your
tongue. And lo and behold, the stone stuck. It was indeed a dinosaur bone! I
was thrilled. I immediately set about searching the surrounding area for
more pieces. I found a second one and then several more. In the end, I had
five elements that all seemed to belong to the same bone. It was too big to
be a turtle bone and too thick and too long to be that of a crocodile, but at
the same time much too small to be the bone of a long-necked dinosaur. The
only animal that made sense for it to have belonged to, in my opinion, was a
carnivore. Judging by the size of the bone, the dinosaur seemed to have
been quite large. I suspected that the fragments were parts of the fibula,
because they appeared too thin for a tibia. I was pleased with this unusual
find, since such carnivore remains are much rarer than those of herbivores.
Since they appeared at the top of a hill, they could not have been washed
there by the rain or any other water movement, and must have weathered
out of the rock in place. When fossils are not transported by wind or water
after they are deposited, they are called “in situ” finds. As excited as I was,
I unfortunately didn’t have a shovel with me to dig deeper into the rock. I
had no choice but to take the exact GPS coordinates, photograph the site,
and then with a heavy heart hike back to the off-road vehicles with the
fragments.
 I proudly showed Oliver my find in the evening and he confirmed that it
was indeed the bone of a larger predatory dinosaur. Oliver had described a
carnivore from the same sediments in 2017, which he had named
Pandoravenator. My fossil was of similar size and was probably another
specimen of Pandoravenator, and perhaps even a new species.
Unfortunately, we could not identify it with certainty because we had to
leave the next day and end the campaign. Another excavation with the
appropriate equipment will hopefully reveal more material and then perhaps
tell us what species it belongs to.

OceanofPDF.com
 Chapter 6

Myanmar—Trapped in “Liquid Gold”

Feathers, Claws, and Ticks in Amber

Michael Crichton’s cleverest idea in his 1990 novel Jurassic Park was to
clone dinosaurs using DNA retrieved from their blood, found inside of
mosquitoes. Crichton wrote that mosquitoes drank the blood of dinosaurs
and immediately became encased in tree sap, which turned to amber over
millions of years. Researchers then extracted the dinosaur blood, decoded
the DNA, and cloned the dinosaur. This approach is currently impossible,
yet the idea fascinates me to this day. There are fantastic amber deposits
from the time of the dinosaurs, and some of them contain inclusions of
spiders, snails, millipedes, and insects, some of which are actually
mosquitoes. The oldest mosquitoes date back to the Cretaceous, but we
have yet to see them from the Jurassic or Triassic. Consequently, a park of
cloned dinosaurs could have included animals like Tyrannosaurus,
Triceratops, raptors, and pterosaurs like Pteranodon, but not Brachiosaurus
and Stegosaurus. Why the novel was nevertheless called Jurassic Park and
not Cretaceous Park was probably due to the euphony of the name rather
than the fact that the dinosaurs were really mostly Jurassic. In fact, most of
the dinosaurs in the book and the first movie came from the Cretaceous
period. Furthermore, in the Jurassic World trilogy, which hit theaters as a
sequel to the Jurassic Park film series, even marine reptiles like the
Mosasaurus made an appearance. Now, that doesn’t make any sense at all
—how would a mosquito have been able to suck blood from this sea
creature, and how could amber have been formed in the sea?
The most important amber deposits with inclusions from the Cretaceous
period come from Canada and Myanmar. The fossils hidden in the amber of
Canada are about 75 million years old and those from Myanmar about 99
million years. Because the country of Myanmar used to be called Burma, it
is still often referred to as Burmese amber or Burmite.
Crichton’s idea of extracting blood from mosquitoes was ingenious, but
he could not have known that about thirty years later the first tiny dinosaurs
would be found in the amber. These finds are truly breathtaking! Burmese
amber gives us detailed information about what the plumage, soft tissues,
and bone structure of juvenile birdlike dinosaurs were like, which we would
not find under any other conditions.
What also makes amber fossils so special is their three-dimensional
preservation. In the Solnhofen Limestone or in shale deposits, animals are
usually flattened, so their actual anatomy is distorted. However, amber
pieces are often small and rarely contain complete vertebrates, but usually
only wingtips or the tail end of very small creatures.
One inclusion described in 2017 contained a very well-preserved
hatchling of a birdlike dinosaur belonging to the enantiornithine
(Enantiornithes) group. This group of feathered dinosaurs is very closely
related to birds, even though it is outside the group of modern birds. Their
strange name means “the opposite birds” and refers to the fact that these
animals were the closest relatives of modern birds and resembled them in
many aspects. However, these enantiornithines are now extinct.
The amber fossil in question holds a just-hatched juvenile that was only a
few hours to a few days old at most. This is indicated by measurements of
the skull, the wing, and the foot, which were compared with other juvenile
and adult enantiornithines. The proportions of the fossil resemble those of
an embryo rather than juveniles. The skull is larger, and the flight feathers
are smaller. In addition, the small size suggests that it is a neonate. Because
of the unique soft tissue preservation, for the first time, it was possible to
observe in a dinosaur the external opening of the ear, the eyelids and scales
on the legs, and the long talons on the toes, with one of the toes facing the
others, so that the animal was probably able to hold on to branches well.
The specimen also offers glimpses of the hatchling’s down feathers, as well
as the arrangement of its pennaceous feathers and even the color pattern of
the plumage. This plumage has characteristics of both precociality and
altriciality at the same time, which is not found in any modern bird. Thus, in
the fossil we find strong flight feathers, while the rest of the body is
sparsely feathered. The feathers on the legs, feet, and tail resemble the
contour feathers of modern birds rather than their down feathers. We also
find additional filamentous, or hairlike, feathers in the same places, which
are comparable to the protofeathers of more primitive theropods. Overall,
these amber finds provide us with a level of detail that fundamentally
improves our understanding of the anatomy of the young of these pre-
modern birds. In the process, it shows that the plumage of enantiornithines
differs markedly from that of modern birds.
Although the above-mentioned animal was a hatchling, it apparently
already possessed rectrices (flight feathers at the end of the tail, designed
for steering), suggesting that it was capable of flight immediately after
hatching. This is consistent with observations in other finds of
enantiornithine juveniles, which are also thought to have been precocial (the
young of precocial birds are already mature and mobile from the moment of
hatching). Extant birds have a wide developmental range in their offspring,
from helpless nestlings born blind and naked to extremely precocious
fledglings able to fly as early as one day after hatching. In contrast, the
range appears to be much narrower among enantiornithines, with every
indication that their chicks were basically precocious. This certainly has to
do with the limited variability in their egg size—all eggs of enantiornithines
are about the same size. Limited developmental variability would also
likely have prevented enantiornithines from breeding in some of the
extreme environments used by extant birds today, where chicks would
otherwise perish without intensive parental care. One example of adaptation
to an extreme ecosystem is that of penguins, whose chicks cannot survive in
the perpetual ice without their parents; another is the adaptation of birds to
live on or in the water, where they must be either good swimmers or good
gliders. So far, there is no evidence for either adaptation in the
enantiornithines. We now know hundreds of aquatic birds, but so far, no
enantiornithines with aquatic adaptations have been discovered. Very
broadly speaking, it can be said that eggs of birds differ drastically in shape,
color, and size depending on whether they breed in trees, cliffs, brushes, or
in the sand. The degree of parental care in enantiornithines is not well
understood today, but apparently their chicks were all highly precocial and
arboreal at the same time. That is a combination we do not observe in any
birds living today. All of today’s tree-dwelling birds have chicks that are
altricial. This means that the enantiornithines followed a developmental
strategy that does not occur in modern birds, as the relatively early sexual
maturity in the young, their slow growth rate, and their slow adolescence
are atypical of birds today. There is a good reason for this: namely, the
retardation in growth meant that these young enantiornithines were longer
exposed to the danger of predation, which increased their mortality rate—
especially if the parents could not protect their chicks because they had left
the nest early. This is why we find so many juveniles of enantiornithines in
the fossil record and virtually none of modern birds.
The chicks from the Myanmar amber allow us to draw conclusions about
the feather development of these avian relatives. When we find certain
types of feathers that are no longer found in adults, or when we see feathers
in adults that juveniles do not yet have, we can see which feathers emerged
only in the course of their ontogeny and how they grew. This is especially
relevant with feathers that we no longer find in modern birds. We often see
such unusual feathers on the legs and the tail. There, a small number of
protofeathers have been preserved, which were part of the down plumage
and were lost in the course of evolution. The legs, the tail, and the belly of
this enantiornithine were sparsely covered with such down, showing that it
had hatched only recently. His molt thus followed the pattern of modern
birds.
Not only are the feathers themselves exciting, but also the ticks that are
sometimes found on them. In 2017, such animals were found on the feather
of a dinosaur. Ticks are now among the most widespread blood-feeding
parasites that live on the skin of a host. For a long time, nothing was known
about the feeding habits and possible hosts of Cretaceous ticks. The feather
found in 2017 now provided direct evidence that ticks fed on the blood of
feathered dinosaurs 99 million years ago. Unfortunately, what kind of
dinosaur it was is still unclear. Because we only have one isolated feather,
the host could have been an enantiornithine or even a bird. Sickle-clawed
raptors or egg thieves are also possibilities. So, in Jurassic Park, John
Hammond, the director of the dinosaur park, could have been looking for
not only mosquitoes but also ticks in amber from which to clone his
dinosaurs.

A Dinosaur Even Smaller than a Hummingbird?

In the competition for the most newly described dinosaurs, Othniel Charles
Marsh and Edward Drinker Cope left behind a hopeless mess of invalid
names and duplicate descriptions of these animals. This had to do with,
among other things, the extreme ambition and egocentricity of the two
paleontologists. Each wanted to outdo the other. Yet, both having been
financially independent, they would have had enough time to examine
specimens more thoroughly and diligently, to proceed with care and
conscientiousness. After all, the number of scientific publications was low
at that time, and the two were able to publish their reports without the need
for lengthy proofreading. Both scientists were wealthy and not necessarily
dependent on research funds—at least not at the beginning of their careers.
Consequently, apart from their own ambition, there was no reason for them
to rush out results. Today, things are different. Research moves so fast that
hundreds of articles are published and up to fifty new dinosaur species are
scientifically described every year.

Scientific articles must be proofread by at least two

independent editors and checked for accuracy of content and
plausibility. Only when the reviewers approve the article may
it be printed in a scientific journal. This process is called peer
review.

At the same time, scientists today are competing for research funding.
Not every researcher receives sufficient funding to advance their field and
publish their findings. There is often not enough money for new equipment,
new computers, to attend meetings, or go into the field. A research proposal
must be written for each grant, and the rejection rate is often high. This is
especially frustrating for young academics. Those who want to stay in
research must show universities that they are intellectually capable of
writing research proposals and performing at a high level. Among the most
important publications for paleontologists are the journals Science and
Nature. For example, to publish an article in Nature, the topic must be
groundbreaking, and the research must be pioneering and involve new
methods. It must appeal to people beyond specialist circles.
The fact that what I perceive as one of the biggest blunders in
paleontology in recent years happened in the renowned journal Nature, of
all places, is extremely surprising. In 2020, a Nature article was published
that had everything you would expect from a paper in this journal.
Researchers had shortly before described a tiny reptile encased in amber.
The find was a sensation, as it was said to be an exceptionally well-
preserved birdlike skull. The skull appeared to belong to the smallest known
dinosaur of the Mesozoic era, which probably weighed only a few grams
during its lifetime, and competed with the bee hummingbird for the place of
smallest dinosaur of all time. However, there was a catch to the story. The
fact that spectacular articles have to be published ever more quickly today
may also have led to a hasty publication in this case. What exactly
happened and how did this mistake come about?
The researchers found a lump of amber containing the head of a small
animal, and the rest of the body was missing. This head had large eye
sockets, a round, domed skullcap, and a long pointed beak. When I saw
photos of this fossil, I too immediately thought it was a bird.
Prior to this animal, only seven specimens of enantiornithine birds had
been scientifically described from amber, which included several articulated
skeletal elements. Six of these were juveniles and one was an almost fully
grown adult specimen. They were all smaller than the fossils that came
from typical sedimentary deposits. But this animal was even smaller.
Because it was also thought to be a bird with large eyes and jaws that still
bore teeth, the researchers named it Oculudentavis, which means “eye-
toothed bird.” The animal was fascinating. In addition to dwarfing and
avian features, it had a unique eye morphology that resembled that of
lizards. In its eye socket was a ring of individual bones called a sclerotic
ring, which indicated that the animal’s pupil had originally been very small,
although the eye socket, and thus probably the eyeball, had been very large.
This bony sclerotic ring apparently indicated a reinforcement of the eye. It
occurs in birds, in the extinct dinosaurs, pterosaurs, and ichthyosaurs, and
also in some modern reptiles. The way these ossicles are arranged is more
reminiscent of squamates than birds or dinosaurs. In every known dinosaur,
the sclerotic ossicles are square shaped and narrow, and they usually delimit
a larger opening inside the ring. In Oculudentavis, the sclerotic ring was
very large, leaving only room for a small pupil. It was formed by elongated,
spoon-shaped ossicles, otherwise found only in lizards.
Deep-diving ichthyosaurs had the largest eyes of any vertebrate and
particularly pronounced sclerotic rings to withstand the enormous water
pressure at the deepest depths of the sea. In the small pupils of
Oculudentavis, my colleagues see evidence of a diurnal lifestyle. Its tiny
body size suggests island dwarfing—according to them, the amber forest
therefore once grew on an island. The unusual body shape of the animal
suggests that this species occupied a previously unknown ecological niche.
With the help of such amber deposits, we are able to uncover habitats that
were inhabited by vertebrate species at the lowest limit of the body size
spectrum. From the tip of the beak to the occiput, the head of this presumed
dinosaur measures only 7.1 millimeters. Except for hummingbirds, there is
no other group of birds that have such a small skull. The skull of
Oculudentavis showed a general avian morphology with a slender beak that
narrows at the tip, nasal openings located far back on its beak, enlarged and
well-defined eye sockets, a short region behind the eye, and a dome-shaped
skull roof. What should have puzzled colleagues, however, were the
unusual teeth. They are stuck in the jaw in a way usually seen only in
iguanas, monitor lizards, and geckos, and not in dinosaurs. Paleontologists
assumed that this unusual feature was related to dwarfism, although no
other detail, such as a shortened beak or toothlessness, indicated
paedomorphosis. Oculudentavis had an entire row of small teeth. Therefore,
the researchers believed that the dwarfing had been accomplished simply by
reducing the growth rate. The way in which the individual bones of the
skull were fused together and the shape of the sutures on the contact
surfaces of the bones also did not conform to the pattern of theropod
dinosaurs, which only emphasized the enigmatic nature of Oculudentavis.
The researchers also tried to attribute this to structural constraints due to
dwarfism. In the end, however, the differences in cranial anatomy were too
great to be explained by processes of insular dwarfism. This was also
noticed by many readers after the article appeared in Nature, which
eventually led the authors to retract it.
Then, in August 2021, a new article on the subject appeared, clearly
showing that the supposed dinosaur was in fact not a bird at all, but a small
lizard. The large eyes and the strongly domed occiput had initially led the
researchers down the wrong track. But a more thorough examination
showed that several features disqualified the animal as a dinosaur despite its
birdlike appearance. Another nearly complete fossil of this prehistoric
animal showed that Oculudentavis was a squamate. The group of squamates
includes nearly all extant reptiles today: lizards, iguanas, monitor lizards,
and snakes. The only modern reptiles that do not belong to squamates are
the turtles, the crocodiles, and the tuatara. An accurate classification in the
animal kingdom had proven so difficult because Oculudentavis had
apparently undergone a convergent evolution of its skull proportions that
made it look like a bird. However, other body features showed a clear
affiliation with the squamates. When skeletal features are weighted and
combined with features from the gene database of extant lizards,
Oculudentavis appears to be closely related to an armless and legless group
of Southeast Asian blind skinks (Dibamidae). If, however, skeletal features
are weighted less, these odd animals end up closer to tuatara in the
phylogenetic tree. If only skeletal features are considered, Oculudentavis
lies somewhere between iguanas and a group of extinct marine reptiles
known as mosasaurs, which could grow up to fifty-nine feet long. With
these animals, Oculudentavis shares a long process of the premaxillary bone
that extends to the nose and the far-recessed nasal openings, but it differs
from them in most other features of the skull. It is all very complex. Despite
this, or perhaps because of it, this fossil is so extremely important to
paleontology. Oculudentavis was part of a forest ecosystem that is
otherwise very rarely preserved in the fossil record, because forests are full
of scavengers, and the forest floor is just teeming with insects and worms
cleaning up the mess. What is not eaten by scavengers and worms is then
taken by fungi. When an animal dies in the forest, there is rarely anything
left of it. The forest recycles very thoroughly and is therefore a bad place
for fossil preservation. The only way for such small organisms to withstand
the millions of years and be preserved as fossils is to be sealed airtight in
tree sap. Amber is therefore a window to the Earth’s past, giving us
glimpses of a fauna that would never have been preserved in sandstone or
mudstone. The conditions of deposition thus dictate what we find later in
the fossil record. In the fossilized tree resin, it is small animals, plants, and
pollen, whereas in the sedimentary rock, it is large bones and tree trunks.
Fortunately, the amber deposits from Myanmar provide us with
remarkable details about the smallest vertebrates of the Mesozoic and help
us in their classification. For example, the very next specimen of
Oculudentavis discovered showed us that although bird features were
present in the skull of the animal, there was no convergence with birds in
the rest of the skeleton. On closer inspection, the resemblance to a bird skull
could be attributed to deformation of the bones during fossilization, but
because the rest of the body was missing from the first specimen, this
misinterpretation was not immediately apparent. We now understand the
ecology of Oculudentavis much better. Its small pupils indeed suggest that
the animal was diurnal, and its embedding in amber suggests that it lived in
trees. The long narrow mandible, the sharp pointed teeth, and the
mandibular joint with little room for jaw muscles and an awkward lever
indicate a weak bite force and a snapping mode of feeding for the animal,
which apparently preyed on insects. The large eyes helped it to better
identify its small prey. The fact that the relationships of Oculudentavis
could be clarified was related to the fact that a second find contained parts
of its body that helped identify the animal as a squamate. In addition, with
the help of a computer program, the researchers were able to virtually re-
create the original morphology of its deformed bones.

When bones are compressed, crushed or shifted against each

other during the fossilization process, this is called
“deformation.” When a computer reverses this process in a
simulation, it is called “retrodeformation.”

Retrodeformation restored the original appearance of the bones, which

made them easier to compare with modern birds and lizards. The
retrodeformation method supports traditional osteological studies, which
will also help in the description of other fossils in the future. New finds
from the amber of Myanmar will hopefully provide more specimens of
Oculudentavis. In particular, finds with bones of its pelvic region, its tail,
and its hands and feet would be important to determine an accurate
phylogenetic position of these bizarre animals.
However, it is not only the discovery of new fossils in Burmese amber
that poses a challenge to scientists; the extremely tense political situation in
the country is also of great concern to my colleagues.

A Moral Dilemma
Fossils from the amber of Myanmar are spectacular because they preserve
structures that usually have a low preservation potential and cannot be
found anywhere else in the world. We can learn a lot about extinct species
from this, because no fossil embedded in a rock matrix provides us with a
comparably detailed picture of the world 99 million years ago. It becomes
increasingly clear that the fossils of the faunal assemblage originated from
an island and contained different species than other fossil deposits from the
mainland. Perhaps someday it will indeed be possible to extract organic
molecules, proteins, or even DNA from dinosaurs in amber, be it through
blood ingested by trapped mosquitoes or directly through the cells, blood
vessels, proteins, or other tissues of small dinosaurs. This prospect is
tantalizing and promises groundbreaking insights in the future.
The amber from Myanmar, however, always plunges paleontologists into
a moral dilemma. Myanmar is one of the poorest countries in the world and
has been politically unstable for decades. Since the military coup on
February 1, 2021, the situation there has deteriorated considerably.
However, as the export of amber strengthens the illegitimate regime, many
of my colleagues are discussing the reconcilability of scientific research
with human rights violations. The Society of Vertebrate Paleontology, or
SVP, a nonprofit scientific organization and the largest association of
researchers, students, and interested laypersons in my field of research, has
contacted its more than two thousand members several times to call
attention to this human rights violation. The society, whose goal is to
advance the science of vertebrate paleontology and to promote the
discovery, preservation, and protection of vertebrate fossils and fossil
deposits, first proposed a publication stop on all scientific articles dealing
with fossils in Myanmar amber recovered after January 2021.
However, the troubling political developments in Myanmar came to a
head when the country’s armed forces seized political control in February
2021 through said military coup and declared a state of emergency. In the
process, over seven hundred people were killed in just two months. The
military had already gained control of the amber mines since 2017,
repeatedly instigated armed conflicts, and carried out ethnic cleansings. The
United Nations condemned these actions as genocide and crimes against
humanity. Since the gemstone mines and amber deposits are located in the
north of the country, where it is very difficult for international observers to
reach, it was easier for the military government to commit atrocities there
unobserved by media representatives and unpunished by the world. They
secured access to the valuable mines and acted with great brutality against
the local rural population. It was important to the SVP that its members and
other paleontologists take special care in their research on the amber
material from Myanmar. They wanted to make sure that fossils did not
come from holdings of the military dictatorship so as not to indirectly
support those in power in the country through research funds. The society
admonished all scientists and scholars to refrain from publishing
manuscripts on Myanmar amber acquired after the 2021 coup. However, the
proportion of amber containing important vertebrate fossils is vanishingly
small, while the majority of amber is processed in the jewelry industry.
Nevertheless, it was important to take a stand on this issue. Even if one’s
own contribution does not necessarily lead to a change in the situation, it is
still important to condemn the injustice and not be complicit in it. The
fossils from this area of conflict are incredibly significant for our
understanding of the evolution of the smallest vertebrates from the
Cretaceous. We can only hope that peace will return to this beautiful
country and that new spectacular fossil finds can be made in the amber that
will reveal to us further secrets from a sunken world.

OceanofPDF.com
 Chapter 7

The Spinosaurs

The Adventures of Ernst Stromer in Egypt

While Werner Janensch was uncovering giant long-necked dinosaurs in
German East Africa, another German paleontologist and important dinosaur
researcher was on the move in North Africa: Karl Heinrich Ernst Freiherr
Stromer von Reichenbach. Stromer, who came from a wealthy noble family
in Nuremberg, began his studies in geology and paleontology at the
University of Munich in 1893, and wrote his dissertation on the geology of
the German colonies in Africa as a doctoral student of Karl Alfred von
Zittel. Africa fascinated Stromer throughout his life, and he came to Egypt
for the first time after his habilitation in the winter of 1901. His first trip
took him to El Fayum, a fossil site discovered by German botanist Georg
August Schweinfurth, which yielded mammals from the Eocene and
Oligocene. Supported by the Bavarian Academy of Sciences, Stromer
conducted a second expedition to Egypt in the winter of 1903/1904 and
finally a third in the winter of 1910. This third expedition was to become
the most important of his career. He arrived in Alexandria in early
November 1910, first to search for early mammals and then to explore the
Bahariya area in the White Desert, which consisted of Cretaceous
sedimentary rocks. However, due to global political tensions on the eve of
World War I, it was difficult for a German explorer to get permission for
such an expedition from English, French, and Egyptian authorities.
However, because of his aristocratic status, financial resources, and social
influence, Stromer eventually succeeded in obtaining the necessary permits.
He met in November 1910 with the British geologist John Ball, who had
just created a geographic map of Egypt and was in the process of publishing
a geological map of the country. Ball gave Stromer both maps, which were
invaluable to the paleontologist. After all, the area around Bahariya was
hardly known at that time, and an expedition into the western desert meant
a dangerous adventure. For example, it was always necessary to find new
grazing grounds for the camels, since the local workers had not bought
fodder for the animals in advance, despite an agreement.
After more than a week of hiking, Stromer’s team finally arrived at the
Bahariya Oasis. But because of the inaccuracy of the geological map, he
could not correctly assess the nature of the rocks around Bahariya, and
Stromer mistakenly believed that the oasis dated to the Eocene, not the
Cretaceous. In mid-January, weather conditions improved, and he began to
explore the Gebel el Dist area. On the very first day, Stromer found a fossil
shark vertebra, fish teeth, and some petrified wood, and four days later, he
discovered three additional large bones that he wanted to excavate and
photograph. The first bone was a femur, badly weathered and incomplete,
but 43.3 inches long and 5.9 inches thick. A second and better-preserved
femur lay next to it; it was 37.4 inches long and as thick as the other. The
third bone was stuck deep in the ground and could not be recovered at first.
After that, Stromer discovered the pelvic bone of a dinosaur, several
vertebrae, and a gigantic claw. In all, he found the remains of four very
large, previously unknown dinosaurs in the Bahariya Basin. Three of them
were carnivores: Bahariasaurus, Carcharodontosaurus, and the famous
Spinosaurus aegyptiacus. The fourth animal was a sauropod, which he later
named Aegyptosaurus.
In addition to dinosaurs, Ernst Stromer and Austrian fossil hunter Richard
Markgraf also excavated snakes, turtles, marine reptiles, and crocodiles
there. However, since the excavation team had not brought any plaster,
reinforcement fabric, or linen, Stromer had to improvise. Out of necessity,
he cut up his mosquito net and soaked it in a mixture of flour and water to
encase the two larger bones with it. The next day, he moved the expedition
to Gebel Umm Hammad, where he also found several dinosaur, fish, and
shark remains, and set out again two days later.
In February 1911 Stromer returned to Germany and over the next few
years studied his finds. In 1912 he described Spinosaurus aegyptiacus and
in 1931 another large carnivore, Carcharodontosaurus saharicus, which
means “the shark lizard from the Sahara.” Stromer chose this name because
the animal’s teeth had serrated cutting edges, reminding him of those of a
great white shark, whose scientific name is Carcharodon carcharias. In
1932 followed the description of Aegyptosaurus (“the Egyptian lizard”) and
finally in 1934 that of Bahariasaurus (“the lizard from Bahariya”).
Until 1914, excavations were still taking place around the Bahariya
Oasis; then World War I broke out, and all the fossils that Stromer had
found were held in Egypt. It was not until long after the end of the war, in
1922, that these finds reached Germany. They were badly damaged because
they had been inspected with little care by the colonial authorities and
stored improperly. The fossils probably would have been held in Cairo even
longer if a Swiss paleontologist, a friend and former student of Stromer’s,
had not taken care of the paperwork and transported the fossils to Germany.
Because of Switzerland’s neutrality, this friend was able to talk to the
British and French and get the pieces out of the country. Fortunately,
Spinosaurus, Stromer’s most important discovery, was not affected, as he
had already taken it with him on his return trip from Egypt. The name
Spinosaurus means “the spine lizard” and refers to the small, winglike
projections of bone that point outward from each vertebra along the spine,
also known as spinous processes. This animal was one of the largest
predatory dinosaurs ever discovered, and we assume that the unusually long
spines on its back likely had skin connecting them, forming a sail-like
structure.
After the National Socialists rose to power in the 1930s, Stromer refused
to join the NSDAP. He stood up for a Jewish colleague, incurring the
displeasure of the party. When World War II began, Stromer’s sons were
drafted, and two of them fell on the battlefield. The third became a Russian
prisoner of war and did not return home until five years after the war ended.
Stromer tried in vain to convince the young and party-loyal curator of the
Munich Museum of Natural History to remove the dinosaurs he had
discovered in Egypt from the collection and bring them to safety. But the
man refused because Stromer had publicly refused to swear allegiance to
the Führer and he assumed out of arrogance that there would be no air raid
on Munich. However, when the Royal Air Force bombed the museum on
April 24, 1944, Stromer’s collections were burned. The skeletons of
Spinosaurus, Carcharodontosaurus, and Aegyptosaurus were completely
destroyed. Ernst Stromer, whose life’s work had been destroyed by the war,
died a broken man in 1952. His work received little attention for several
decades. Only when a new paleontological expedition to Bahariya was
undertaken in 2000, and the remains of more dinosaurs were recovered
there, did his research come back into the limelight. Among the new finds
was the sauropod Paralititan stromeri, which bears Stromer’s name in its
honor. Whether the interest in his work also contributed to the appearance
of Spinosaurus in the third Jurassic Park film, I cannot say. But I am sure
that the paleontological advisors of the movie read the scientific description
of the Paralititan and thought it would be cool to have the Tyrannosaurus
and the Spinosaurus fight each other.
The question of how big and how heavy Spinosaurus actually was has
long been the subject of great disagreement among experts. So far, no
complete skeleton has been found, and the nature of its anatomy is hotly
disputed. Most recent estimates, however, assume a length of about fifty
feet, and there are hardly any deviations from this anymore. But as far as
his weight is concerned, there is still great uncertainty. Depending on which
calculations one follows, the animal is supposed to have weighed between
four and up to twenty tons. More recent estimates are between six and
seven and a half tons.
I am asked surprisingly often who would win in a real fight, the
Tyrannosaurus or the Spinosaurus. I assume that this question is related to a
scene from the third Jurassic Park film from 2001, in which a Spinosaurus
effortlessly kills a Tyrannosaurus. Despite the fact that a time span of about
30 million years separates the two animals geologically, and that they
would never have met geographically (Tyrannosaurus lived in North
America and Spinosaurus in the Mediterranean), there is probably no doubt
about the outcome of such a fight. Spinosaurus with its fifty feet was
certainly somewhat longer than the approximately forty-foot-long
Tyrannosaurus. However, the length gives only limited information about
the mass of an animal. The extinct giant snake Titanoboa was probably
much longer than most tyrannosaurs, but I don’t think that a constrictor
snake would have stood a chance against the king of the dinosaurs with his
enormous teeth. It is similar with Spinosaurus. Its teeth—unlike those of
most other carnivorous dinosaurs—were round in cross-section and were
used more for snatching and holding prey than for cutting meat, biting
tendons, or breaking and crushing bones and carapaces. The differences are
striking. Tyrannosaurus teeth were longer, rooted deeper in the jaw,
sturdier, and had cutting edges to bite huge chunks of flesh out of their
victims. But the most obvious difference is the power with which a
Tyrannosaurus could bite. Calculations have shown that it had the strongest
bite force of any land creature ever. The bite force of a Tyrannosaurus was
about three times as powerful as that of a Spinosaurus. And while
Spinosaurus’s prey consisted of smaller or medium-sized animals it picked
up while wading through lakes and rivers, Tyrannosaurus was capable of
hunting and killing large herbivores, some of which were armored. Still, the
two would probably have preferred to stay out of each other’s way because,
of course, there would still have been a risk of injury to Tyrannosaurus in a
fight. However, if I had to make a bet on the outcome of such a fight, I
would always bet on Tyrannosaurus.

Spinosaurus: On Land, on the Shore,

or in the Water?

Spinosaurus is one of the most popular and well-known dinosaurs, and ever
since new remains of it were discovered in the Kem Kem Basin in
Morocco, researchers around the world have been taking a closer look at it.
And because it’s such an unusual dinosaur, distinctly different from other
carnivores, discussions about its appearance, lifestyle, diet, size, and weight
continue to inspire strange hypotheses. But I’ve been upset with the public
perception of Spinosaurus and its portrayal in the media lately.
It all started with an article by Nizar Ibrahim and his colleagues in 2014
that described Spinosaurus as semiaquatic. There is a lot to be said for
Spinosaurus living near water. It certainly stayed on the shores of streams
and lakes, and its diet consisted to a not insignificant extent of fish. This is
supported by the shape of its teeth, which resemble those of crocodiles and
marine reptiles, and also by the shape of its skull, which is more like that of
a crocodile than those of other carnivorous dinosaurs. In the article written
by Ibrahim and other scientists, the team of authors attempted to prove an
aquatic lifestyle by highlighting the fleshy nostrils located far back on the
head, and the unusually long neck and torso for a carnivore, which shift the
animal’s center of mass well forward of its pelvis. In addition, the authors
point out the short hind legs, which on land would have led to a
disadvantage in hunting. Probably their most important argument is that the
leg bones did not have an open medullary cavity and instead had a massive
bony cortex on the shaft. Such bones are indeed indicative of a secondary
adaptation to life in the sea, as we observe today in penguins, for example,
which have very massive bones compared to other birds. If you look at the
ribs in manatees and dugongs, you will notice that they are additionally
broadened and thick and also lack a medullary cavity.

Secondary marine animals show such extra-wide ribs or thick

limb bones without medullary cavity, one speaks of
pachyostotic bones or pachyostosis. In my example, it is a
secondary adaptation of manatees and dugongs to a life in
water, acquired through evolution. In order to compensate for
the static buoyancy of the body and to be able to remain
effortlessly underwater, the average density of the animal
increases, while the thickness of the bones increases, and the
circumference of the medullary cavity decreases at the same
time. The bones thus perform a function similar to a diver’s
belt.

The team of authors succeeded in making a big splash on the topic; their
article appeared in the prestigious journal Science, while National
Geographic simultaneously published a long popular science article about
their research. An Italian company made a life-size model of a Spinosaurus
that was featured on the cover of National Geographic, and a Canadian
company built a museum-quality skeletal model. Ibrahim gave a perfectly
staged talk at the largest meeting of vertebrate paleontologists, and a movie
was even made about how he serendipitously ran into a fossil dealer in
Morocco who led him to a secret site in the Kem Kem Desert where new
remains of Spinosaurus were found. The story was a huge media spectacle,
and I seemed to be the only one who didn’t take the bait. This was back
when I was still working at the dinosaur park, where they were also in the
process of making a new model of Spinosaurus. I remember well the
discussions I had with the park’s general manager, the sculptor, the
scientific director, and a former fellow student from Bonn who was now
also working in the park. They all found Nizar Ibrahim’s article interesting
and wanted to build the model according to his specifications. My
criticisms fell on deaf ears.
A heated exchange with the managing director, in which he pointed out to
me that four votes were in favor of these changes and only one vote (mine)
was against them, was the last straw. I replied undiplomatically that science
is not a democracy and only the facts count. After that, it was clear that I
would not be working in the park much longer.
What bothered me was the fact that Ibrahim’s reconstruction was based
on a composite of several individuals, whose proportions I thought he had
wrongly scaled and misinterpreted. The animal looked kind of jumbled
together even at first glance.
The second argument that made no sense to me was the reference to the
pachyostotic bones. I had already seen, during excursions to the Mainz
Basin, fossils of sea cows that had swam around with their thick, broad ribs
in the shallow sea there during the Oligocene, about 30 million years ago;
moreover, I had observed pachyostosis on a primitive mammal, the
enigmatic Desmostylus in the collection of the Tsukuba University in Japan,
and therefore I knew that the bones of Spinosaurus were quite different in
nature.
Apart from that, it was absolutely incomprehensible to me why the leg
bones in an animal should be thicker, but at the same time shorter, to give it
additional weight. On the contrary, this animal would have to become rather
lighter by such a development, because the mass gain by pachyostotic
bones would not have outweighed the mass loss of smaller muscles on the
smaller legs.
Also, I did not see any significant widening of the ribs, which would
have been much more important for static lift. And why, of all things, would
an animal with such a massive dorsal sail live in the water? A crocodile
ambushing unsuspecting wildebeest or zebra at a water trough might be
completely submerged until only its nostrils peek out of the water. In the
same situation, a spinosaur’s dorsal sail would have protruded out of the
water by up to six feet. This would certainly not have been a good strategy
for an ambush predator, whose hunting success heavily depended on the
surprise effect. If Spinosaurus had then also decided to swim or submerge
so far that its sail would also have been underwater, the water resistance
would have made any change of direction impossible, and the animal would
have drifted helplessly as soon as the sail accidentally got crosswise to the
direction of flow.
Ibrahim and his colleagues only marginally addressed the sail and its
hydrodynamic properties. Instead, they argued that this body part was
probably a display feature males used to make an impression on females. I
don’t doubt that, because on land such an imposing structure would
certainly have caused a stir, but in the water, it would only have been a
hindrance.
I don’t blame the media here—the whole story was sensational and
exciting, because Spinosaurus would have been the first non-avian dinosaur
adapted to a life in water. Dinosaurs with such a high degree of adaptation
to water are only found among today’s birds. Penguins are the only birds
that show quite extraordinary adaptations to life in the sea.
To test Ibrahim’s hypotheses, Donald Henderson created three-
dimensional digital models of Spinosaurus and other animals in 2018 that
also accounted for regional density variations in the animals’ lungs and air
sacs. In doing so, Henderson noted that the model of Spinosaurus tended to
tip on its side in the water. His models of Baryonyx, Tyrannosaurus,
Allosaurus, Struthiomimus, and Coelophysis showed similar buoyancy,
suggesting that Spinosaurus’s hydrodynamic properties were far from
exceptional. The software also provided evidence that the center of mass of
Spinosaurus was close to the hip. This would allow Spinosaurus to move
forward well on land. Its pneumatized skeleton and air sacs ensured that it
was unsinkable. A semiaquatic lifestyle of pursuit hunting in the water is
thus ruled out. Henderson concluded that Spinosaurus may have been more
specialized for a life on the shoreline, wading only occasionally in shallow
water. At the same time, it may have been a competent hunter on land,
spending a significant portion of its life away from the water.
The article about the static buoyancy of this dinosaur did not receive
nearly as much attention as the media spectacle about the swimming
Spinosaurus. After Henderson’s article appeared, I pointed out his new
findings in an online forum and also mentioned the unfavorable
hydrodynamic properties of the large dorsal sail. I must admit that this was
somewhat naive, as my contribution was immediately criticized, and I was
personally insulted and responded to with hostility. I don’t normally
converse in these forums, so I was unfamiliar with the strange dynamics of
such groups, where anyone can just post anything without backing up their
claims with appropriate scientific evidence. I was asked who I was that I
could dare to doubt the “fact” of an aquatic lifestyle of Spinosaurus. When I
looked at the profile of my vociferous critic, to my knowledge this person
had no academic background, nor did he practice any scientific profession.
Though this topic is heavily debated and there are many opposing
theories, I don’t understand at all why everyone seems to want to make
Spinosaurus into a creature that it is not. With its massive size, huge dorsal
sail, and crocodile-like snout, Spinosaurus is one of the most fascinating
dinosaurs, whether it was aquatic or not, and its popularity today almost
rivals that of Tyrannosaurus. Unfortunately, the movie Jurassic Park III and
the 2014 publication previously mentioned have contributed to the public’s
misconception of Spinosaurus. It was such an extraordinary animal that
there is no need to attribute additional superpowers to it. In 2020, Nizar
Ibrahim published a new study to debunk Henderson’s arguments and
provide more evidence of strong aquatic adaptation. In principle, I think
highly of maintaining discourse on a controversial topic when new evidence
emerges. However, sometimes one has to wonder if new work actually adds
to our scientific understanding or has been published for questionable
reasons. While Ibrahim and his team acknowledged that Henderson had
presented compelling anatomical, biomechanical, and taphonomic reasons
to question a semiaquatic lifestyle, they also presented evidence for a
putative swimming tail. Their article appeared this time in the prestigious
journal Nature. They argued that Spinosaurus possessed extremely high
dorsal and ventral spinous processes on its caudal vertebrae, forming a large
and flexible finlike organ with which it could supposedly perform wide-
ranging swimming movements. To demonstrate these swimming
characteristics, they built several underwater models of oscillating tails,
measured the physical forces on them, and compared different tail shapes.
They wanted to show that the tail of Spinosaurus produced greater thrust in
the water than that of other supposedly land-dwelling dinosaurs, and that
Spinosaurus was therefore more comparable to aquatic vertebrates, which
had laterally flattened and vertically extended swimming tails to produce
propulsion while swimming. Unfortunately, the story was again
inconclusive from my perspective. Namely, the team claimed that all
Spinosaurus relatives had the characteristic crocodile-like teeth and
elongated snout, suggesting that they were all piscivores, yet the tails of the
remaining spinosaurids showed only minor aquatic adaptations.
 What bothered me about this experimental setup was that the two-
dimensional robotic models of the swimming tails could only inaccurately
simulate the movements of a three-dimensional body in the water. This was
also noticed by David Hone, a colleague from London, and Thomas Holtz,
one of the world’s leading experts on predatory dinosaurs, who took a
closer look at Ibrahim’s study. They included new finds of Spinosaurus in
their considerations. The two paleontologists evaluated arguments about the
animal’s functional morphology, ecology, and its hunting and feeding habits
in the context of these new finds. They were convinced that the degree of
Ibrahim’s aquatic adaptations was overestimated. The interpretation of the
anatomy of the tail as a caudal fin was not tenable in their view, and they
expressed considerable doubt that Spinosaurus was able to swim efficiently
and rapidly with this tail and to pursue prey underwater. They saw no
evidence for a highly specialized adaptation as an aquatic predator and
instead proposed a model that Spinosaurus fished predominantly along the
coast or in shallow waters but could also search inland for prey. Even
evidence that Spinosaurus was a good swimmer would not be
counterevidence for a wading lifestyle and the fact that this dinosaur may
have sought its food along the coast. Conversely, the instability in the water,
the high water resistance, the position of the eyes and the nostrils, and the
low swimming performance argued against an efficient hunter underwater.
Another aspect that had been noticed in previous studies was the underside
of its neck, which had rugose muscle attachments, indicating strong
musculature that may have been useful for rapid, downward pecking
movements of the head. The animal may have fished in the water with its
snout, as storks do today. It may have remained underwater with its mouth
slightly open until a fish swam by, and then snapped at it, or lunged at the
prey with a rapid downward movement. The ability for such rapid
downward movements would not have benefited a pursuit predator
underwater because there, fish can escape in all directions. In this context,
the position of its nostrils far back on the skull is advantageous, since
Spinosaurus could still breathe easily even with its mouth underwater, as
storks can. David Hone and Thomas Holtz therefore drew a picture of a
generalist that searched for prey along the shore, in and out of the water,
and also did not disdain meat of animals that had come to their death at
watering holes or during floods. The denser leg bones would have aided its
wading because they would have minimized unwanted buoyancy in the
water. By being able to hunt in the water and on land, there was less
competition for these animals from crocodiles and land-only predators, and
they could also move more easily from one lake to the next.
There is another piece of evidence that supports this hypothesis.
Examinations of an animal’s tooth enamel can show what it feeds on
predominantly. Certain isotopes reveal whether it ate more land animals or
fish. In the case of Spinosaurus, this signal is ambiguous because it
probably made its prey both on land and in the water.
In late 2022, Professor Paul Sereno of Chicago joined the discussion
about Spinosaurus. Ibrahim was Sereno’s student and Sereno had been one
of the coauthors of his 2014 article. Therefore, I was surprised when Sereno
criticized Ibrahim’s second Spinosaurus article from Nature and came to a
completely different conclusion. Paul Sereno and his colleagues used
computed tomography to scan all available parts from different specimens
of the animal and created a three-dimensional model to which they added
muscles, air sacs, and lungs to model the weight distribution as realistically
as possible. They showed that Spinosaurus was only about forty-five feet
long and, while on land, walked bipedally rather than on all fours, while its
body was laterally unstable in deep water. It was a slow surface swimmer
with weak tail muscles that could not move forward in water faster than 2.2
miles per hour. Said paleontologists also demonstrated that Spinosaurus’s
static buoyancy was so great that it certainly could not dive. This dinosaur
could have waded to a water depth of about 8.5 feet but would then have
floated on the surface of the water, unable to submerge. In fact, while its
hind legs had slightly thicker bone walls, its dorsal vertebrae had air-filled
cavities in front of the pelvis, and the bones of its forearms had a
pronounced medullary cavity. Sereno and his colleagues calculated that
because of the strong pneumatization of the vertebrae, Spinosaurus, despite
its considerable weight of about 7.4 metric tons, had a low density of only
about fifty-three pounds per cubic foot, which would have been far too little
to dive on its own.

Salt water has a density of about sixty-four pounds per cubic

foot.
 Sereno’s team pointed out that reptiles living today with similar dorsal
sails or high, laterally flattened tails, such as the common basilisk
(Basiliscus basiliscus), use them for display rather than actual aquatic
locomotion. Vertebrates secondarily adapted to aquatic life, on the other
hand, almost invariably have a fleshy fluke or caudal fin rather than a
flattened swimming tail, and also have greatly reduced limb length.
Spinosaurus, with its powerful legs, on the other hand, could reach far
inland without using its long-clawed hands for weight support. Although its
physique suggests a semiaquatic lifestyle, the research team ruled out a
purely aquatic lifestyle because at no time in the fossil record were there
secondary aquatic vertebrates that lived in fresh water and were longer than
seven feet. Such animals all lived in the open ocean, far from the coast. A
semiaquatic lifestyle, on the other hand, can occur at any body size.

Fish and No Chips:

The Fish Eaters of the Isle of Wight

Ernst Stromer discovered the first remains of Spinosaurus in North Africa,

but only recently have we learned how diverse the spinosaur group is,
through finds on an island in the English Channel: the Isle of Wight. In
2010, a paleontological field trip from the University of Bonn under
Professor Sander took me there for the first time. I immediately fell in love
with the island because of its beauty and stunning dinosaur fossils. You can
only reach it by ferry from Portsmouth or Lymington, which dock in the
north of the island. However, dinosaurs and their footprints are found on the
Isle of Wight only in the southwest and southeast. The rocks in which their
fossils are found belong to the so-called Wealden Group; they date from the
Lower Cretaceous and are about 125 to 140 million years old.
 At the south coast of the Isle of Wight, United Kingdom, natural casts of dinosaurian footprints are
exposed.
PHOTO BY THE AUTHOR

The south coast of the Isle of Wight is very steep and experiences strong
erosion. There, fossilized footprints of dinosaurs crop out, which are
gradually washed into the sea. If they are not salvaged, they will eventually
be swallowed up by the waves. Such footprints were formed when a
dinosaur walked on the soft clay or sandy ground and sank a bit into the
subsoil, leaving imprints. These impression molds were then infilled with
sand or clay shortly after their formation and were thus protected. This
allowed both the soil and the infillings, called casts, to slowly turn into
stone. Sometimes the rock that forms these casts is harder or more resistant
to weathering than the surrounding matrix, and when the matrix weathers
away, millions of years later, only the casts remain. Sometimes these casts
are not quite clearly identifiable as such, but if you have an eye for it you
can easily spot them. On the Isle of Wight, footprints of dinosaurs are found
far more frequently than their bones. A single imprint in sedimentary rock is
simply called a footprint, while several footprints in succession are a track,
and the layer on which such a track is found is a track site.
 The most common tracks found on the Isle of Wight are those of
iguanodontids. Iguanodon was one of the largest ornithischian dinosaurs of
the Lower Cretaceous. After Megalosaurus, Iguanodon was the second
dinosaur to be scientifically described, in 1825. Particularly striking are its
bony thumbs, which were initially misinterpreted as a nasal horn. The first
reconstructions show Iguanodon with a quadrupedal, or four-legged, mode
of locomotion. Later, it was depicted as bipedal for nearly a hundred years.
But more recently it is assumed that it actually walked mainly on all fours
and only moved bipedally when it fled from danger. Tracks of Iguanodon
confirm this assumption. Since its hoof-like forefeet were much smaller, it
is easy to distinguish impressions of their hands from their feet. Another
herbivorous ornithischian dinosaur found on the island is Hypsilophodon. It
was significantly smaller and actually a nimble biped.

Iguanodon and Hypsilophodon possessed three toes, the outer two of

which were splayed at a ninety-degree angle. Predatory dinosaurs have very
similar tracks to these two animals, but their toes are usually more slender,
and the outer toes form an angle of less than ninety degrees. The
impressions of predatory dinosaurs on a track site sometimes also retain the
hollow shapes of the sharp claws, which facilitate assignment. In Germany,
for example, such footprints can be found at the Dinosaur Park
Münchehagen. The sediments in which they occur are as old as those on the
Isle of Wight. The special thing about the dinosaurs on the English island,
however, is not only the high number of finds, but above all the incredibly
diverse dinosaurian fauna. Even long-necked dinosaurs such as Cetiosaurus
and Ornithopsis were found there. A particularly complete skeleton of a
long-necked dinosaur is the so-called Barnes High sauropod. In 1992,
almost its entire skeleton, including the head, was found, which is very rare
in sauropods. The animal still has no scientific name, but it was closely
related to Brachiosaurus and is considered the most complete sauropod
skeleton of England. There are also remains of armored dinosaurs on the
Isle of Wight, such as Polacanthus, whose name means “many thorns.” It is
an early representative of the ankylosaurs that carried spines along their
flanks and bony plates—called osteoderms—on their backs. These
osteoderms were fused together over the pelvis to form some kind of armor.
With all these finds, it is not surprising that the Isle of Wight is also called
the Dinosaur Island of England. More than twenty-five different species of
dinosaurs are already known from there and that doesn’t even include the
undescribed finds. When these species are also eventually described, the
number will increase significantly. What makes the dinosaur fauna of this
island so interesting for research is the especially high number and diversity
of predatory dinosaurs.

The Needles form the western tip of the Isle of Wight and are the main tourist attraction of the island.
PHOTO BY THE AUTHOR
 With a body length of up to twenty-two feet, Neovenator was the largest
purely carnivorous dinosaur on the Isle of Wight. It could move quickly and
had three deadly claws on each foot and razor-sharp teeth in its mouth.
Unlike Tyrannosaurus, however, these teeth were very thin and better for
cutting meat than for crushing or biting bone. Tyrannosaurs also play a role
in the fauna of the island. In fact, the oldest representative of the group and
thus the earliest ancestor of Tyrannosaurus rex was discovered on the Isle
of Wight. Its name is Eotyrannus, which means “the tyrant of the dawn.”
He was about sixteen feet long and was described only in this century. Even
bigger than Neovenator was Baryonyx. It was probably not a pure
carnivore, but rather a fish-eater, as suggested by its teeth and its crocodile-
like skull shape. Baryonyx was a close relative of Spinosaurus but did not
have a dorsal sail. Its skeleton is the most complete of any spinosaur to
date. In 2021, two new spinosaurids were described: Riparovenator—“the
hunter of the riverbank”—and Ceratosuchops, “the horned crocodile face.”
However, not many remains of these two animals have been found. In 2022,
another spinosaurid was reported. It too appears to be a new species, as its
fossils were found in younger strata, the Vectis Formation on the southwest
coast of the island, near the village of Compton Chine, which—strictly
speaking—consists of only a farm and a large parking lot. The find contains
no skull bones, which is why researchers have not yet assigned a scientific
name to the animal. Cervical, pelvic, and caudal vertebrae are well
preserved, as well as parts of the ilium, ribs, and fragments of arm or leg
bones. These remains suggest an individual of enormous proportions,
comparable to the size of the Spinosaurus found by Stromer. This would
make the animal much larger than the other known spinosaurs from the
island. And, assuming similar anatomy to Spinosaurus, this find could even
be the largest carnivorous dinosaur ever found in Europe. A comparison
with the bones of other spinosaurids shows that this yet unnamed animal is
more closely related to Spinosaurus than to the other spinosaurids of the
Isle of Wight, such as Baryonyx, Ceratosuchops, and Riparovenator. The
find from the Vectis Formation also suggests that spinosaurids diversified
and occupied new ecological niches since their origin. While the earlier
representatives from the Wessex Formation predominantly inhabited the
banks of extensive river systems, this animal apparently lived near a lagoon
from which few other dinosaurs are known to date.
 What fascinates me is that spinosaurids probably always occurred in
ecosystems that had very high carnivore diversity. We see that with Ernst
Stromer’s dinosaurs from the Bahariya Oasis, which included Spinosaurus,
Carcharodontosaurus, and Bahariasaurus. And the same is true on the Isle
of Wight, where, besides the three spinosaurids Baryonyx walkeri,
Ceratosuchops inferodios, and Riparovenator milnerae, there may have
been a fourth, much larger genus from the same family, plus the
tyrannosauroid Eotyrannus lengi, the large predator Neovenator salerii, and
a number of other carnivores whose position in the dinosaur family tree is
not entirely clear.
In Morocco we have a similar picture. There lived besides Spinosaurus
also Carcharodontosaurus and huge crocodiles, which were so big that they
probably ate even dinosaurs. And on the Isle of Wight, the fragmentary
remains of the new spinosaurid suggest that it too may have been one of the
largest carnivores in Europe.
We will learn later, in the chapter on Tyrannosaurus, that this was very
unusual for dinosaurian faunas. And the fact that in these assemblages oft
particularly large predators occurred, makes little sense in regard to the
distribution of large predators within the trophic network.

OceanofPDF.com
 Chapter 8

More than Just Bones

The Super-Lung
Through the various fossil discoveries, we have learned much about the
powerful lungs of dinosaurs. We see evidence of unidirectional lungs in the
highly pneumatized bones of the vertebrae, and we can infer their
functioning from the anatomy of modern birds and crocodiles. Many of
these conclusions are based on computer models and theoretical
considerations. Yet for a long time there was no clear, direct evidence
because the lung, like all other soft tissues, has a low preservation potential.
But that changed when an animal called Archaeorhynchus was reported
from China that was already more closely related to modern birds than to
enantiornithines. It came from the Jiufotang Formation of the Lower
Cretaceous, and its soft tissue and plumage were still very well preserved.
In its thorax were the remains of two lungs that resembled those of living
birds, an indication that even these early forms of birds were capable of
flight 120 million years ago. Of all living, air-breathing vertebrates, birds
have the most complex and efficient respiratory system, allowing them to
maintain their energetically demanding form of locomotion even in oxygen-
deficient environments. The lung microstructure of Archaeorhynchus also
appears modern and is further evidence that many physiological changes in
the respiratory system that characterize extant birds and helped them
succeed were already present in dinosaurs that were not directly on the
avian lineage. Instead of saying that dinosaurs possessed birdlike lungs, it
would probably be more accurate to say that birds possess dinosaur lungs.
 The Cloaca:
How Do Dinosaurs Make Wee-Wee?

Walking through the halls of the Senckenberg Museum in Frankfurt, it’s

easy to lose track of all the treasures the museum has to offer. In the large
hall, skeletons of a long-necked dinosaur, a horned dinosaur, an
iguanodontid, a club-tail dinosaur, and a tyrannosaur are on display. Behind
it lies the famous dinosaur mummy. In this setting, this original fossil of a
small herbivore hardly catches the eye, lying inconspicuously near the wall
in a display case. It is the Frankfurt specimen of a Psittacosaurus from the
Early Cretaceous Jehol deposits of Liaoning in China. This fossil has the
best preservation of scale-covered skin of any ornithischian dinosaur
described to date. One can even discern its color pattern and
countershading, which allows us to reconstruct in detail the appearance of
this animal and draw conclusions about its habitat. The countershading
suggests that the animal lived in a shaded, forested environment. Because
the skin and scales are so well preserved, this piece also allows us to see a
cloaca on a dinosaur for the first time.
The Psittacosaurus specimen of the Senckenberg Museum in Frankfurt, Germany, shows remarkable
soft tissue preservation.
PHOTO BY THE AUTHOR

A cloaca is the posterior orifice that serves as the only

opening for the digestive, reproductive, and urinary tracts. It
was originally present in all vertebrates but was replaced by
separate excretory orifices in bony fish and most placental
mammals. The cloaca is the posterior section of the rectum, to
which the reproductive organs and the ureters are connected.
Through it, sperm, ova, and excrement are discharged via the
anus.

The fact that dinosaurs had a cloaca is basically no surprise, because it is

also true for all archosaurs living today, even if it looks different in birds
and crocodiles.
 To better compare the small dinosaur with its living relatives, the
specimen containing the fossil was scanned, which allowed
retrodeformation of its virtual model. The preservation of the rectum of this
Psittacosaurus is remarkably good and shows in detail the anatomy of its
cloaca. Its opening is clearly visible and can be readily distinguished from
surrounding tissues, as the shape of the scales and pigmentation differs from
those of adjacent body regions. Although the cloaca gives no indication of
the sex of the animal, the pigments appear to have been signal colors,
possibly intended to create a visual stimulus. One might think of it as in
baboons or mandrills, whose rumps are colored differently than the rest of
their fur.

Do Female Dinosaurs Have Ovaries?

Hard-shelled eggs are a characteristic shared by all birds and crocodiles. Yet
the reproductive systems of all living archosaurs are very different—as are
their cloacae. Crocodiles have two ovaries, their ovarian follicles mature
slowly, and they have large clutches and small eggs. Living birds, on the
other hand, almost all have only one ovary and one oviduct, and their
follicles develop rapidly. Birds are now the most diverse tetrapod class, and
there is a wide range of clutch and egg sizes within their group, depending
on the body size of the bird. Generally, however, the eggs of birds are
larger, and their clutches are smaller than those of crocodilians.
Derived reproductive traits in birds are thought to have evolved gradually
early in the dinosaur evolution, but the exact timing of these changes is
difficult to trace because of the incompleteness of the fossil record and
because of sparse soft tissue finds. However, we have learned much from
fantastic finds from the Jehol Province in northeastern China. Recently,
there have been reports of exceptionally well-preserved ovarian follicles in
the prehistoric bird Jeholornis and in several enantiornithines from the
Early Cretaceous strata of this region. Initially, it was doubted whether
these were actually follicles or rather seeds that had been eaten by the
animals.
To find out, fragments of these alleged follicles were extracted and
subjected to several analyses. These structures were found to have
histological and histochemical characteristics of smooth muscle fibers
intertwined with collagen fibers, resembling the contractile structure of
connective tissue in birds today. Fossilized blood vessels were also
preserved. No plant fibers or evidence for other plant tissue was found,
supporting the original interpretation as follicles in the left ovary. At the
same time, the right ovary was absent, showing that it was apparently
functionally lost early in avian evolution. The fossils from China were
compared with the small carnivore Compsognathus from Germany, which,
like Archaeopteryx, was found in the Solnhofen Limestone.
As it turned out, Jeholornis and Compsognathus both show features
similar to those of present-day crocodiles because they still have a large
number of small ovarian follicles. Enantiornithines also differ from modern
birds in terms of ovarian follicles. They had fewer and larger follicles;
however, these follicles were all similar in size. In today’s chickens, the size
of the follicles in the ovary varies, corresponding to different stages of
development.

The Grace of the Sleeping Club-Tail Dragon

Kai Jäger, one of my former fellow students from Bonn, dealt in his
research with the evolution of early mammals. Kai can talk about his work
in a very lively and humorous way, thus making it more accessible to a
broad audience. That’s why he also took part in Science Slam, a
competition in which researchers have to present their results as
entertainingly as possible in a very short period of time. The focus is on
communicating scientific content in a way that is broadly accessible. The
participants’ ten-minute presentations are voted on by the audience through
cheering with the winner being the one who receives the most applause. Kai
did so well that he was invited to the German Science Slam Championship
in Berlin, which he actually won in 2014. Since then, he has been regularly
invited to speak by radio and television broadcasters when an expert in
paleontology is needed. Hence, it was not surprising when in 2017 the
Deutschlandfunk radio station called him when a spectacular find of a club-
tail dinosaur was reported from Canada. However, he forwarded this
request to me, since he is not a real dinosaur expert, but as a paleontologist
is mainly interested in the evolution of mammals. This radio program was
about an ankylosaur that had been discovered in the oil sands of Canada,
and since I had already read quite a bit about it, I was familiar with the
topic. The animal, named Borealopelta markmitchelli, was found in a mine
in northeastern Alberta. There, workers came across its skeleton in a debris
heap, and employees of the mining company immediately informed the
Royal Tyrrell Museum in Drumheller, one of the best-known and most
famous paleontological museums in the world. It has one of the largest and
most speciose collections of dinosaurs, with an emphasis on fossil finds
from the Canadian Badlands. A museum team traveled directly to this mine,
recovered the fossil with help from the workers, and shipped it back to
Drumheller. The rock strata in which the animal was found is composed of
Albian marine sediments and belongs to the Clearwater Formation, where
several plesiosaurs and ichthyosaurs were also deposited. However, no
dinosaur had ever been discovered in it until that time, which is not
surprising because the formation contains sediments of a coastal facies and
a nearshore marine environment. Normally, land creatures would not be
present in these sediments. Most likely, said Borealopelta had fallen into a
river and had been washed away by it. Whether this happened before or
after the death of the animal is not conclusively clear. In any case, after its
death, the carcass, floating on its back, entered the open water and then
sank to the bottom of the sea. There, the animal was quickly buried by
sediment. Although remains of seafloor benthos such as worms, snails, and
starfish can be found in these deposits, there was no evidence of feeding
marks on the dinosaur by such scavengers. If, despite the seafloor being
oxygen-rich and populated by burrowing animals, the carcass does not
show any feeding marks, it is an unmistakable sign of a very rapid burial.
When the fossil was found, it was completely encased in a dense and hard
—but at the same time brittle—iron carbonate concretion, such as the one
we have already encountered with the ichthyosaurs from Nevada. Cracks on
the surface of the concretion showed that it must have fractured as the
carcass collapsed in its calcareous grave, as the organic material
increasingly decomposed, liquefied, and produced decomposition gases.
When the loading pressure finally became too great, the gases escaped, and
the body fluids were released. The resulting cavity in the dinosaur’s carcass
was then filled with sediment. However, because the animal was lying
underwater with its belly side up, its carapace remained intact. As a result,
not only have its scales and osteoderms been almost completely preserved,
but so have their arrangement in their natural compound, and—much more
unusual—many of the dermal bone plates are still covered by a horny
sheath. Because the empty body cavity was so quickly filled with sand, the
fossil is barely crushed, and we now have an excellent three-dimensional
preservation. The Canadian research team was even able to identify the
pattern of its scales and systematically searched for melanin pigments
which were preserved on the carapace.

Melanin are widespread dark brown to black or sometimes

yellowish or reddish pigments that cause coloration of the
skin, hair, feathers, or eyes in animals.

To reveal all of this, the fossil had to be freed from the concretion in the
laboratory first, which took the preparator Mark Mitchell a full five years
and around seven thousand hours of work. His extraordinary achievement
was recognized with the naming of the animal, whose species name ended
up being markmitchelli. The genus name Borealopelta is a combination of
Latin and Greek and means “the shield from the north,” alluding to its
northern location and its carapace. Mark Mitchell’s effort was clearly worth
it, because the fossil is very beautiful. For its display, they turned it upside
down so that the carapace was again facing up. If you look at it from the
front, you might think Borealopelta is still alive and just sleeping, which
has earned it the nickname “Sleeping Beauty.” Although the animal may not
be as pretty as a king’s daughter, at least it looks like a sleeping dragon,
with its entire front part preserved. The head, neck, most of the torso, and
its limbs were protected by osteoderms during its lifetime. On its flanks, the
animal also bore long spines. One might think it was a walking tank on four
legs. But the color pigments of its carapace reveal that even adult
Borealopelta must have had predators. The pigments are distributed over its
carapace in a countershading pattern and were meant to fool predators.

When the coloration of an animal is darker on the upper side

and lighter on the lower side of the body, it is called
countershading. Because this phenomenon was first observed
and described by the painter Abbott Thayer, it is also called
the Thayer’s principle. Countershading is a form of
camouflage because it makes it more difficult for predators to
see their prey.

Countershading was already observable in ichthyosaurs, and we will

encounter this survival strategy again in other dinosaurs in a later chapter.
When I was asked about this in the radio interview on Deutschlandfunk, I
expressed doubts about the interpretation of the carapace coloration of
Borealopelta. I was of the opinion that Borealopelta did not need
camouflage because it was simply too large and too well armored. Even
large tortoises, where small members of the group still have camouflage
colors, lose their coloration without predation pressure. With an estimated
length of eighteen feet and a body mass of at least twenty-nine hundred
pounds, Borealopelta was much larger than any modern land animal that
exhibits countershading. In today’s mammalian faunas, predators pose no
risk to herbivores weighing more than a ton. These huge animals do not
need to hide and therefore have no camouflage. If they also have defensive
weapons such as horns or tusks, predation pressure decreases even further.
So why would Borealopelta have needed camouflage? I found it more
plausible that its reddish coloration represented a form of aposematism
(advertising signals in the form of conspicuous coloration) that some
venomous animals display. I could also imagine that the strong coloration
might have played a role in mating and courtship. However, I had not
considered that the predator-prey situation was completely different in the
Cretaceous. While there are no remains of carnivores in the same strata, we
do know of footprints of particularly large predators in the Clearwater
Formation from descendants of allosaurs. Weighing several tons, these
theropods were strong enough to kill even an adult club-tail dinosaur; they
were more than ten times heavier than the largest predators of the present
day (such as tigers, lions, and polar bears). We’ll learn more about the
difference between dinosaur-and mammal-dominated ecosystems when we
get to tyrannosaurs. But the hunters of Borealopelta were not
tyrannosaurids. In fact, they didn’t appear until about 50 million years later.
The fact that camouflage was especially important in dinosaurs,
crocodiles, and birds is apparently also related to the exceptionally good
eyesight of modern birds and crocodiles. We assume that these groups
inherited the ability from their common ancestors, and therefore assume
that the large theropods also had very good vision and that these carnivores
were very visual predators. This is in contrast to modern ecosystems, where
large mammals at the top of the food chain can only perceive a limited
spectrum of colors, even when compared to humans. The enormous size
and strong visual dependence of Cretaceous apex predators may have led to
an evolutionary arms race that resulted in a combination of armor and
camouflage even in the largest herbivorous dinosaurs. Thus, club-tail
dinosaurs also had to hide and camouflage. The presence of countershading
in a large, heavily armored herbivorous dinosaur thus offers us a unique
glimpse into a Cretaceous predator-prey dynamic that is not comparable to
processes in nature today.

OceanofPDF.com
The Upper Cretaceous

(100.5 to 66 million years

before present)

OceanofPDF.com
OceanofPDF.com
 Chapter 9

Wyoming—The Hell Creek

Triceratops: A Tank on Four Legs

One of the most important paleontologists of our time, Professor Roger
Benson, once said that there was one correct answer to the question of
which dinosaur was the best: Triceratops!
That was just a joke, of course, but I was pleased anyway, because
Triceratops has always been my favorite dinosaur. Even in my childhood, I
enthusiastically gazed at the skulls of Triceratops in Frankfurt’s
Senckenberg Museum. My poor brother unfortunately had to be there with
me, too, although he was not at all a dinosaur fan. Over the years I’ve seen
many skeletons, but I always like to return to Frankfurt to take a selfie with
Triceratops. The animal is on display in Frankfurt, although it is actually
from Wyoming. Triceratops, along with Tyrannosaurus rex, was one of the
last dinosaurs outside the avian lineage. Remains of Triceratops and
Tyrannosaurus are found in the same strata. They come from the Lance
Formation and the Hell Creek Formation, which were deposited during the
last 2 million years of the Cretaceous period.
 The two skulls of Triceratops, discovered by Charles Sternberg, now on display at the Senckenberg
Museum in Frankfurt, Germany.
PHOTO BY THE AUTHOR

Interest in Triceratops skeletons has been high in recent years, with

several auctions of these fossils causing a stir. In October 2021, one
particularly massive specimen sold for 6.65 million euros (about 7.7 million
US dollars at the time) to a private collector at an auction in Paris. At the
time, people worried that the animal would no longer be accessible to the
public and to researchers in the future. Fortunately, however, the fossil
found its way to the museum of the D’Annunzio University of Chieti–
Pescara in Italy. The giant skeleton was discovered in South Dakota in 2014
in the Hell Creek Formation. It is about 23.5 feet long and has a hip height
of almost nine feet. Because of its size, it is also nicknamed Big John. The
skull alone accounts for more than one third of the animal’s total length. It
is the largest nearly complete Triceratops skeleton in the world; 60 percent
of the body and over 75 percent of the skull have been preserved. However,
we know of partial skeletons and isolated skulls that suggest there must
have been much larger specimens. The word triceratops means “three-
horned face.” This dinosaur had two long horns over its eyes and one on its
nose. It belongs to the group Certatopsia, or horned dinosaurs. Almost
everyone knows the name Triceratops, but surprisingly few people know
that this dinosaur, just like Tyrannosaurus rex and every other scientifically
described animal, also has a species name. There are two valid species
names of Triceratops: Triceratops horridus and Triceratops prorsus.
Horridus (Latin for “rough” or “wrinkled”) refers to the uneven surface of
its bones and prorsus (Latin for “forward”) refers to the forward-pointing
nasal horn. A total of seventeen different species of Triceratops have been
described, with all but the two mentioned now invalid. When horns of a
Triceratops were first discovered, they were thought to be the horns of an
extinct species of bison and the find was named Bison alticornis, meaning
“bison with high horns.” If you count this name, the Triceratops has
eighteen species names.

The two valid names of Triceratops: T. prorsus, T. horridus

Invalid names used as synonyms for the valid species: T.
albertensis, T. alticornis (originally Bison alticornis), T.
brevicornus, T. calicornis, T. elatus, T. eurycephalus, T.
flabellatus, T. galeus, T. hatcheri, T. ingens, T. maximus, T.
mortuarius, T. obtusus, T. serratus, T. sulcatus, T. sylvestris.

Triceratops was first described by Othniel Charles Marsh in 1889. The

horns, which he thought were those of a bison, came from the Denver
region of Colorado. The first whole skeleton came from the Lance
Formation in Wyoming. Triceratops bones were also found in the Hell
Creek Formation. This dinosaur was hunted by the most dangerous predator
of all time, Tyrannosaurus rex. But Triceratops was also an extremely
strong and dangerous animal and by no means defenseless. Tyrannosaurus
could only attack by ambushing Triceratops. It is quite possible that some
tyrannosaurs did not survive their attack on a Triceratops because they
themselves were seriously injured in the process. The bony, three-foot-long
horns above the eyes of Triceratops were dangerous weapons. Like those of
cows and buffalo, they were covered with a horn sheath made of keratin.
The animal had a thick, massive frill that protected its delicate neck,
shoulders, and nape. Triceratops’s head was connected to its spine by a
ball-and-socket joint, so that it could turn in all directions at lightning speed
and an attacker always had to stare at its pointed horns first. If Triceratops
was not caught by surprise, it was very difficult to kill it. The first real
evidence of a duel between a Tyrannosaurus and a Triceratops was
provided by a spectacular fossil discovered in Montana in 2006, which is
now on display in Raleigh, North Carolina. The fossil contains a not-yet-
fully-grown Tyrannosaurus and a Triceratops that are preserved, joined in
mortal combat. The young T. rex obviously overestimated itself in the
selection of its prey, and in the end both adversaries came to death.
Triceratops was rather quarrelsome and frequently engaged in territorial
and mating fights, proven by its injuries on the frill of Big John. It bears a
puncture wound that, judging by its shape and size, was inflicted by the
horn of a conspecific. Although the wound had already partially healed on
the edges, it must have become infected while the animal was still alive.
Perhaps it was so deep that Big John later died from the infection. However,
the neck shield of Triceratops served not only as a defense, but also as a
means of attracting a mate. It is quite possible that these shields had
conspicuous display colors. Some scientists initially even thought that
horned dinosaurs developed their frills to distinguish between different
horned dinosaur species and to separate their own species from others, but
this has since been ruled out. Triceratops had an additional bone in the
lower jaw that is found only in ornithischian dinosaurs such as duckbill
dinosaurs and horned or armored dinosaurs. This bone lies anterior to the
mandible and is called the predental. It is thus a synapomorphy of the
ornithischian dinosaurs.
There still is no consensus among experts on how exactly the
ornithischian dinosaurs developed. It was assumed earlier that Silesaurus,
the small omnivore from the Middle and Upper Triassic of Poland, which
we have already met at the beginning of this book, was maybe one of the
first ornithischian dinosaurs. But this is more than unlikely. The fossils of
Silesaurus come from the Polish Drawno Beds Formation, which formed
about 230 million years ago. In addition to skeletons, fossil footprints called
Atreipus were also found there, and were initially thought to have come
from ornithischian dinosaurs. It is now clear that Silesaurus left these
tracks. The skull of this animal was narrow and, just like that of Triceratops
many millions of years later, bore a horned beak at the tip of the lower jaw
that resembled the predental. However, because the predental is exclusive to
ornithischian dinosaurs, this feature was considered evidence of Silesaurus
belonging to the group of ornithischian dinosaurs. In reality, however,
Silesaurus did not have a predental. Its horned beak evolved convergently.
Triceratops had another bone in the upper jaw, in front of the premaxilla,
which is found exclusively in horned dinosaurs. This so-called rostral was
toothless and looked like a parrot’s beak. It formed the counterpart to the
predental and is a synapomorphy of the horned dinosaurs, which gave them
their typical appearance.

How a Cretaceous Landscape is Resurrected

The two Triceratops skulls in Frankfurt’s Senckenberg Museum are my

favorite fossils in this exhibition. I can still vividly remember standing in
front of the fossils in amazement as a little boy and my father snapping a
photo of my brother and me. I was so excited and fascinated. That was in
the early 1980s. These fossils were offered for sale to the museum as early
as 1910, and that same year it acquired another important dinosaur fossil:
the Trachodon mummy. This is a nearly complete skeleton of a duckbill
dinosaur, now named Edmontosaurus annectens. It is so unusual because it
is still largely covered with its scaly skin. The name Trachodon (“mummy”)
goes back, as it should, to the squabblers Othniel Charles Marsh and
Edward Drinker Cope. An associate of Marsh discovered the first known
remains of the animal in 1891, and Marsh named it Claosaurus annectens.
However, because he had not been very thorough, the animal was renamed
many times in an attempt to undo erroneous descriptions and duplicate
names. It was called Trachodon for a while, and was given the name
Edmontosaurus as recently as 2011. Therefore, Trachodon is now more of a
nickname rather than a valid scientific name.
My brother (left) and me in front of Sternberg’s Triceratops skulls during my family’s first visit to the
Senckenberg Museum.
PHOTO SUPPLIED BY THE AUTHOR
 Charles Sternberg’s Edmontosaurus mummy, now on display at the Senckenberg Museum in
Frankfurt, Germany.
PHOTO BY THE AUTHOR

One of the most talented fossil hunters who worked for both Marsh and
Cope—and probably the most successful—was Charles Hazelius Sternberg.
He accompanied the two paleontologists on various campaigns into the
Cretaceous strata of Wyoming and Montana, and even after the death of the
two disputants in the early twentieth century, Sternberg returned there. It is
thanks to him that Frankfurt has this mummy. Charles Sternberg, who was
sixty years old at the time of its discovery, had previously dug for dinosaurs
in the area for over twenty years and had a forty-year career as a fossil
collector. He had earned his reputation as an expert fossil collector at a
young age when he gathered fossilized plants from near his home in Kansas
for paleobotanist Leo Lesquereux of the Smithsonian Museum in
Washington. Word of his sleuthing soon spread, and a little later, in 1876, he
began searching for dinosaurs full-time for Edward Cope. That was also the
year General Custer was crushingly defeated at the Battle of the Little
Bighorn by the Sioux, Arapaho, and Cheyenne under their leaders Sitting
Bull, Crazy Horse, and Gall in what is now Montana. Not far from this
theater of war, in the middle of the Wild West, Cope and Sternberg
unearthed spectacular dinosaur finds. Marsh also sought good relations with
the Indigenous tribes so that he could dig in their territory. From 1889 to
1894, Sternberg’s son Charles, along with paleontologist John Bell Hatcher,
searched for vertebrate fossils there on Marsh’s behalf and actually found
the first remains of the horned dinosaur Triceratops—even though he
initially mistook its horns as bison horns. Horned dinosaurs are also called
ceratopsians, and because their remains were often found in the strata of the
Lance Formation, Marsh named these deposits “Ceratops beds.” Because of
the constant weathering at the surface of the site, new fossil discoveries
continue to be made there regularly today. According to 2017 and 2018
counts, a total of nearly seven hundred skulls and partial skeletons of
horned dinosaurs, one hundred and fifty skeletons of duckbill dinosaurs,
and about seventy partial tyrannosaur skeletons have been discovered in the
Lance and Hell Creek formations. Teeth and individual bones of dinosaurs
are found even more frequently, with over forty thousand documented from
these strata, private collections excluded.
Cope and Marsh had both been dead for ten years when Sternberg
returned to the region. Starting in 1908, he, together with his sons, George,
Charles Mortam, and Levi, worked in the Lance Formation strata in
Converse County (now Niobrara County), Wyoming. The county was much
larger in the early twentieth century; only later did the eastern part of
Converse County become what is now Niobrara County, and only the
western part retained its old name. In 1908 and 1909, the family discovered
the first Trachodon mummy and two Triceratops skulls. They sold one of
them to the London Museum of Natural History, while the other, along with
the duckbill dinosaur, was acquired by the American Museum of Natural
History. The Senckenberg mummy came from the southern Schneider
Creek area in Niobrara County. In 1910, Charles Mortam, the second-born
son of Charles Sr., discovered parts of a dinosaur tail weathering out of the
sandstone. Subsequent excavation revealed a complete skeleton with skin
impressions. In situ, it measured 17.2 feet, of which the skull accounted for
3.9 feet, the torso 7.9 feet, and the tail 5.4 feet. The recovery of this fossil
was the most laborious that the Sternberg family had undertaken up to that
time. Charles was determined to secure every fragment of the skin
impressions, so the salvaged blocks of rock turned out to be particularly
large. The mummy’s torso alone weighed about 1.6 metric tons, and the
total weight of the fossil was 4.5 metric tons. Since the Sternbergs did not
have a block and tackle, the recovery of the fossil blocks could only be
done step-by-step by lifting them ever so slightly. It is hard to imagine
today how these four men could cope with the tremendous undertaking. The
Sternbergs built a ramp of sand and soil and lifted the fossil out of the pit in
an immense feat of strength. They used poplar logs as levers and then
shoveled sand underneath the blocks of rock. In this way, they were lifted
inch by inch to a height of about 3.9 feet before being hoisted onto the
horse-drawn carriage and taken to the railroad station at Edgemont in South
Dakota, some seventy-five miles away. In total, the excavation took two
and a half months.
Charles Sternberg offered to sell the fossil to Fritz Drevermann, then the
director of the Senckenberg Museum. Drevermann was able to raise the
requested sum through a donation from the industrialist Arthur von
Weinberg. However, shortly after the agreement with the German museum
director, Sternberg received another offer from the Canadian Museum of
Nature in Ottawa. The Canadians offered Charles twice the amount of
money for the fossil that he was to receive from the Senckenberg Museum.
Sternberg wrote about it in his 1917 memoirs: “I shall never forget the
effort I made to induce him to give up the specimen, or take another in its
stead. [...] But [the fossil] crossed the Atlantic. The last message I had of it,
before this awful war [World War I] cut off all communications, was that
the head had been prepared and it was the best of which there was any
record.”
In the summer of 1910, the Sternbergs also discovered four Triceratops
skulls, two of which went to the Senckenberg Museum, and a few years
later Charles Sternberg found another dinosaur mummy, which he sold to
the British Museum of Natural History shortly before the First World War.
The fossil was to be brought to Europe by ship. Unfortunately, that ship
would be the ocean liner RMS Lusitania, which was sunk by a German
submarine not far from the Irish coast on May 7, 1915. This terrible tragedy
claimed the lives of 1,198 people and the sinking of the Lusitania ultimately
led to the United States entering the First World War. This is the reason
why, to this day, the London Museum does not have a dinosaur mummy and
this one lies at the bottom of the sea. More than a hundred years after the
ship first sank, however, there’s probably nothing left of it.
As interesting as the descriptions of fascinating, novel, and ever bigger
dinosaurs may have been back then, nowadays it is no longer enough just to
look for the largest fossils. Today, other research topics have become much
more important. We want to understand the environment the animals lived
in, what they ate, what enemies they had, and what other animals coexisted
in the same ecosystem. Scientists want to find out what climate and
vegetation prevailed at that time, and they want to learn about the ways of
life and cause of death of the dinosaurs. That’s why, in 2019, researchers
from the Senckenberg Museum traveled once again to the original site in
Niobrara County to reconstruct and understand the ecosystem of
Edmontosaurus. In a collaboration with the Wyoming Dinosaur Center
Foundation and supported by the Lipoid Foundation, the Senckenberg team
explored the strata of the Lance Formation and recovered an approximately
215-square-foot bone field in July 2019. I had the honor and the great
pleasure of accompanying the excavation for National Geographic
Germany to write four articles about it. For me, it was a pleasure to be part
of an excavation for the Senckenberg Museum so many years after my first
visit to Frankfurt—even more so in an area where Barnum Brown had
found remains of Tyrannosaurus and Charles Hazelius Sternberg of
Triceratops. So, I took a flight from Frankfurt to Denver and rented a four-
wheel drive SUV to take me to the quiet town of Lusk. It is located in far
eastern Wyoming and has about fifteen hundred inhabitants. That’s 60
percent of the total population of Niobrara County, the most sparsely
populated county in Wyoming. Other than a bank, a supermarket, a truck
stop, and two liquor stores, there is not much in this town, but it is located
in a geologically interesting basin. All around, the land is mostly flat, and
extensive pastures stretch out for the many herds of cattle. Here and there
you see pumping stations that extract petroleum from rock strata a mile
deep. It was here that Sternberg and his sons discovered the Senckenberg
dinosaur mummy, even before the county was established in 1911. The
rocks exposed north of Lusk are part of the Lance Formation, named for the
village of Lance Creek, and contain one of the best-known Late Cretaceous
dinosaur faunas. Anyone even remotely interested in dinosaurs knows the
names of the animals that have been excavated here: Triceratops,
Edmontosaurus, Ankylosaurus, and, of course, Tyrannosaurus.
 The site we drove to was rediscovered back in the 1970s. It is located on
the grounds of a private ranch about fifty miles north of Lusk. To get there,
you exit US Highway 18 at Mule Creek Junction and end up on a gravel
road. Wild sunflowers lined our path, an invasive plant species not
originally endemic to the area. The ride was repeatedly disrupted by cattle
carelessly crossing the gravel road while pronghorn and mule deer grazed in
the distance. With the exception of a small deciduous forest near a river, the
area was treeless for many miles. Just before the gravel road ended after ten
miles, it took us past a colony of prairie dogs that watched each passing car
suspiciously and called out warnings to each other before disappearing into
their burrows. From here, the only way to get farther was with an all-terrain
vehicle with plenty of ground clearance. Although the site had been known
for a long time, it was only then that a scientific excavation was first carried
out, which was certainly related to the fact that such a research project was
expensive and involved considerable effort. In addition, the site was simply
too remote for regular excavations. In the middle of nowhere on the
American prairie, some forty tons of ancient bones and rocks had to be
lifted, cut up, loaded onto a container, and shipped—a logistical challenge
in a class of its own. Fortunately, the Senckenberg team had support at the
time from the Wyoming Dinosaur Center, which had leased the outcrop.
Philipe Havlik, curator at the Senckenberg Museum in Frankfurt, was in
charge of the excavation. For him, the site was important because he and his
science team wanted to understand what had happened there in the Late
Cretaceous. Today, we know that during this time period, there were
particularly high carbon dioxide concentrations in the atmosphere. In order
to find out how this phenomenon affected the vegetation at that time, a
precise profile of the rock sequence was needed. This would provide us
with information about the type of deposition and, in conjunction with
further lab work, enable us to surmise the era’s climate. The scientists at the
Senckenberg Museum were finally able not only to identify numerous plant
and animal fossils, but also to reconstruct the entire ecosystem in which
Edmontosaurus once lived. However, elaborate tooth enamel studies that
provide clues to the chemical composition of the atmosphere at the time,
just like studies of plant pollen, can only be done under ideal conditions in a
scientific laboratory. Even small fossils such as mammal teeth, fish scales,
or plant seeds, which are often overlooked in the field, can only be spotted
and quantified in the laboratory. Hence, the question was how to transfer an
entire bone bed to Frankfurt. Philipe Havlik had an answer: instead of
recovering individual bones as Sternberg once did, he decided on a recovery
en block. We were to cut out all the fossils, along with the surrounding
matrix, as one big chunk of rock. This concept was not new, but to recover a
block of 215 square feet was unique and presented a great challenge even to
the experienced Senckenberg crew. The paleontologist Manuela Aiglstorfer
from Mainz, who accompanied our excavation in the hopes of finding many
fossil mammal remains, explained to me that at first the area had to be
uncovered and several feet of overburden had to be removed with the
excavator in order to reach the fossil-bearing layers. In doing so, it was
important not to damage bones that protruded from the rock. Bones that
were visible on the surface were measured and then either recovered
directly or plastered in place. Long steel bolts were then driven into the
rock layers of the fossil deposit, and ropes were tied to them to create an
excavation grid. This grid helped to mark the exact position of the bones
removed, so that they could be reassigned later, in the lab. A grid was then
drawn with a spray can and cuboids with an edge length of 3.3 feet were cut
out piece by piece. A chain saw with a diamond-tipped saw blade was used
for this work, cooled with water from a large tank. Cutting such a bone bed
into cubes with a saw, when the bone density is so high, one might risk
cutting the bones. But these cuts are rather small compared to the total area
and size of the bones, and one can reassemble the cut bones relatively easily
in the laboratory. After sawing, the cut surfaces of the cuboids were
reinforced with reinforcing fabric soaked in resin to prevent the sediment
from crumbling out. Once a cuboid was suitably prepared, it was removed
by a telehandler, a type of mini-excavator. Steel plates were attached to the
lifting fork, the front edge whetted and its surface greased with lubricating
oil. This allowed the telehandler to use sheer force to push the steel plate
into the in situ rock beneath the cuboid and dislodge it from the subsoil.
Then the oversized “cake lifter” was raised and the remaining surfaces of
the rock were also sealed with resin. Now the cuboid only had to be
shuttered with plywood sheets and lashed down with ratchet straps. Finally,
it was loaded onto a pallet and into an overseas container. Despite all the
auxiliary tools, a lot of manpower was needed to handle such a project. That
is why Philipe Havlik and Manuela Aiglstorfer were supported by a team of
eleven from France.
 As soon as we arrived there, it became clear to us that the effort would be
worth it, because individual bones of the Edmontosaurs were already
sticking out on the surface of the rock. One could see ribs, vertebrae,
humeri, and thigh bones with the naked eye, but also more delicate
elements such as lower jaws and even finger bones. The size of the bones
left no doubt that these were the remains of dinosaurs. Furthermore, the
hooves and teeth were indicative of several duckbill dinosaurs. The way
they were deposited revealed that the bones must have been moved after
death and possibly washed away in a flood, as many different bones from
different individuals were crisscrossed. Interestingly, however, there was
little to no abrasion at the tips of the bones. This suggested a short transport
during which the bones had not rubbed against each other. The exact
number of animals buried here could no longer be determined with absolute
certainty. In such a situation, however, one can use some simple tools to
determine a minimum number of individuals. If, for example, five left
femora are found, one can be certain that there were at least five
individuals, since such a bone occurs only once per animal. If there are
several identical bones of different length, this may speak for animals of
different ontogenetic stages. It was clear that the latter was the case with the
dinosaurs in our bone bed, and also that it was a herd of a single species.
 Close-up of the bone bed in Niobrara County, Wyoming, showing a vertebra (front) and a scapula
(back) of Edmontosaurus.
PHOTO BY THE AUTHOR

Philipe was pleased not only with the dinosaurs, but also with smaller
finds such as soft-shelled turtle shell fragments, the first mammal tooth of
the dig, and another hand-sized piece of rock that showed a deciduous tree
leaf next to a conifer branch. Such a fossil would never be seen in the
Jurassic strata since deciduous trees did not appear until the Cretaceous
period. Layers of leaves or charcoal and individual tree trunks were also
found in the bone bed, repeatedly. There was much to suggest that there
were mixed forests in this region of the Midwest of North America at that
time, indicating to us the prevailing climate at the end of the Cretaceous.
The tension in the field was great because we naturally wanted to get the
bone bed to Frankfurt in one piece. Our schedule turned out to be tight and
the working conditions were difficult, because all the equipment and tools
had to be laboriously transported to the excavation site. In addition, there
was no workshop for miles around and no electricity, water, or sanitary
facilities on-site—only the burning sun and a lack of shade. The extreme
conditions at the excavation site were not only a challenge for the research
team, but also put a strain on the equipment. Each block that had to be cut
weighed about 1.5 tons, and the total weight of the removed sediment was
thirty to forty tons. The chain saw turned out to be the weak point of the
project. There were repeated delays due to its defective metal blade and
chain. Another issue was the crumbling rock, which broke off at the edges
of the blocks despite the resin and could have affected the stability of the
whole block during transport. This was mitigated by superglue, of which
the team used up dozens of bottles during those days. The coat of resin
around the blocks was also a source of danger because, as soon as the toxic,
corrosive compound was mixed, it generated intense heat that could have
caused burns or even caught fire. Therefore, protective gloves were
necessary. Once the resin had been applied and dried, the wooden sheeting
was put in place and fastened with the ratchet straps. Then the cavities were
filled with polyurethane foam to prevent movement within the block during
transport. Whenever a damaged saw blade needed to be replaced, I used the
time to search for more fossils on a nearby hill. Philipe told me that he had
once found small fragments of a Triceratops frill there, and of course I
wanted to see for myself if I might discover more pieces of my favorite
dinosaur. But since I didn’t have the right tools with me, my search was
limited to the terrain surface. At least I also found a few frill fragments.
Overall, this excavation was quite impressive. The rock blocks were
loaded into a large overseas container with the telehandler. This mini-
excavator was just as wide as the inside of the container and was able to
push one pallet at a time deep inside. The overseas container was then
loaded onto a large truck. To this day, it is still a mystery to me how the
truck driver managed to get so close to the excavation site with his huge
truck. Even with our much smaller off-road vehicle, we had to carefully
drive around holes and large boulders on the gravel road so as not to
damage the underbody of the truck. The big truck drove the huge chunk of
dinosaur graveyard two thousand miles to the East Coast. There, the fossil
bones landed on a container ship and, after millions of years underground,
sailed across the vast ocean toward Frankfurt. Later, during the preparation
of the blocks, a few predatory dinosaur teeth were actually found—and they
were from a Tyrannosaurus!
OceanofPDF.com
 Chapter 10

Tyrannosaurus—The Measure of All Things!

The Power of the Brand: Why Tyrannosaurus Is So Popular

Every child today knows Tyrannosaurus rex. It is the most popular and well-
known dinosaur, and is renowned not only in vertebrate paleontology, but
also in the hearts of many children. It is a media phenomenon. No other
dinosaur has been immortalized so many times in film and television. The
scene in Jurassic Park in which the muffled stomp of the Tyrannosaurus can
be heard for the first time, causing the water in the glass to shake even
before the giant is seen, is one of the most iconic and spine-chilling moments
in movie history. Not least because of the enormous success of Jurassic
Park, this animal has become a real merchandising powerhouse. No other
dinosaur is featured more often on T-shirts and bedsheets. It appears on
dinosaur puzzles, on Twitter, YouTube, and TikTok, in memes, cartoons, and
comics, and is certainly the most common dinosaur game character (from
Tomb Raider to World of Warcraft, to Dino Crisis, Dinosaur Hunter, and
many more). It is in every bag of dino nuggets, adorns every child’s birthday
cakes. But what makes Tyrannosaurus so unique? What makes it so popular?
A lot of it has to do with its name: Tyrannosaurus rex! It means “the king of
the tyrant lizards,” and that’s an apt name for the undisputed ruler of the
Cretaceous period. Tyrannosaurus, however, almost had a completely
different name. The first isolated teeth of Tyrannosaurus were found in
Colorado as early as 1874, but at that time they could not be assigned to any
animal. In the 1890s, isolated bones were discovered in eastern Wyoming,
and in 1892, Edward Drinker Cope described two vertebrae of a
Tyrannosaurus, but he mistook them for the vertebrae of a horned dinosaur.
He named the animal to which they belonged Manospondylus gigas,
meaning “large porous vertebrae.” If these vertebrae had been better
preserved, so that the dinosaur could have been distinguished from other
carnivores by their characteristics, then Tyrannosaurus would be called
Manospondylus today. I doubt that this name would have had a similar
appeal. In 1900, the American paleontologist Barnum Brown found the first
partial skeleton of a giant carnivore in eastern Wyoming, where Edward
Cope had also already discovered vertebrae of Manospondylus and the first
complete Triceratops. Two years later, Barnum Brown discovered another
partial skeleton of Tyrannosaurus in the Hell Creek Formation in Montana.
Brown wrote at that time that he had never seen anything like it from the
Cretaceous period. Henry Fairfield Osborn, who in the meantime had been
named president of the American Museum of Natural History in New York,
assigned two names for both animals in a 1905 publication. He called the
find from Wyoming Tyrannosaurus rex and the animal from Montana
Dynamosaurus imperiosus, which means “imperial power lizard.” A year
later, however, he realized that both finds were of the same species, and
fortunately decided on the impressive name Tyrannosaurus. The lower jaw
of the animal he had first named Dynamosaurus can still be seen today at the
Natural History Museum in London. Since Osborn first published both
animal names in the same publication, he could just as easily have chosen
Dynamosaurus. However, Dynamosaurus sounds half as exciting as
Tyrannosaurus, so we can be glad that he chose the latter name, which we
still know and love today! A total of just over thirty complete or at least
nearly complete skeletons are currently known. Two of the most famous
tyrannosaurs were discovered in the 1990s: Stan, long on display at the
Black Hills Institute in South Dakota, and Sue, which can still be admired
today at the Field Museum in Chicago. Stan and Sue take their nicknames
from their respective discoverers, Stan Sacrison and Sue Hendrickson.
One of the most beautiful skeletons of a tyrannosaur is the so-called
Tristan Otto. It was discovered in Montana in 2012 and has the best-
preserved tyrannosaur skull in the world. Tristan was on display at the
Natural History Museum in Berlin from 2015 to 2020, before being
transferred to Copenhagen. It has been back in Berlin since 2022. What
makes Tristan special is its black fossil bones. Minerals give the bones a
silky sheen. In Leiden, Holland, another tyrannosaur skeleton has been on
display since 2016 that is one of the largest and most complete. It also comes
from Montana and is nicknamed Trix. From 2017 to 2019, it was the main
attraction of a traveling exhibition visiting various places in Europe. I saw
Trix in 2018 at a convention in Paris.
 A much smaller specimen of a tyrannosaurid was discovered in 1946.
American paleontologist Robert Bakker and his team gave it the new species
name Nanotryrannus lancensis in 1988, which means “the dwarf tyrant from
the Lance Formation.” It’s a curious name, since the remains actually came
from the Hell Creek Formation. There is disagreement among experts as to
whether this is really a new species or just a young Tyrannosaurus rex.
Recent findings suggest that it was indeed a juvenile of T. rex, but the
relatively long arms and its divergent dental formula cast doubt on this. Why
it is nevertheless plausible, and what allometry and variability have to do
with it, I will come to that later.

A cast of the “large” arms of the juvenile “Nanotyrannus” at the Black Hills Institute, South Dakota.
PHOTO BY THE AUTHOR

One of the most spectacular finds in my opinion was made in Montana in

2006. A Tyrannosaurus and a Triceratops were found together, united in a
death match in one big block. This can be seen as direct evidence that
Tyrannosaurus hunted Triceratops. The fossil was privately owned for
fourteen years until a museum in North Carolina bought it. It was finally put
on display for the public in 2022, and I can’t wait to see it with my own
eyes. The fact that people flock to special exhibits around the world just to
see a real skeleton of these animals shows their unwavering popularity. But
it is not only the public that is interested in Tyrannosaurus. Few other
dinosaurs are as well studied as it is, and in the last five years alone,
paleontologists have discovered more about this animal than ever before.
Today, we understand its biology much better. We know why there were no
other larger predators in ecosystems in which Tyrannosaurus occurred, we
know why its arms were so small, and whether or not it was feathered.
Nevertheless, much is still unexplored, and many mysteries surrounding the
king of dinosaurs have yet to be solved.

A Record Sum for the King

When I was on the dig in the Lance Formation in Wyoming in the summer
of 2019, I met Pete Larson, president of the Black Hills Institute, a dinosaur
museum in Hill City, South Dakota. Pete is an extraordinary personality and
one of the most important Tyrannosaurus discoverers. It is definitely worth
googling his name sometime or watching the movie Dinosaur 13, which
explores the discovery of the tyrannosaur Sue. Pete invited me, the
photographer, and the cameraman from the excavation to visit him in South
Dakota. I really wanted to see his museum because it had Stan, one of the
largest and most complete tyrannosaurs in the world, on display. The animal
is extremely impressive, and I definitely wanted to see it at the Black Hills
Institute before it was sold, as Pete had told us it would be. Pete and his
brother were in litigation at the time, and as a result of the settlement, Pete
would keep the museum while his brother would get the proceeds from
Stan’s sale. Finally, on October 6, 2020, the T. rex was sold to an unknown
bidder for the incredible sum of 31.8 million dollars. This was a record sum
that had never been collected for a fossil before and exceeded the wildest
estimates many times over. The previous record holder at auction had also
been a tyrannosaur, the T. rex nicknamed Sue from the Natural History
Museum in Chicago. Its skeleton had already earned its previous owner a
whopping 8.36 million dollars in 1992. But as great as the enthusiasm for
this high sum was, the experts were shocked. No one knew who the
mysterious bidder was, and it was feared that a private collector or investor
might withdraw the fossil from public view and prevent future research on
the animal. From October 2020 until the end of March 2022, the fossil
remained missing, and speculation about the mysterious owner and the
animal’s whereabouts ran rampant. When a tyrannosaur skull was seen in the
background during a television interview with actor and wrestler Dwayne
“The Rock” Johnson in January 2022, rumors spread that he might be the
one who had purchased the dinosaur at the auction. However, it was only a
replica of Stan’s head. Two months later, it became known that the future
Museum of Natural History in Abu Dhabi had bought the fossil at auction.
Many paleontologists expressed relief that a museum had acquired the
animal and welcomed the fact that it would now be on display in a part of
the world where people rarely had access to fossils and dinosaurs in
particular.

Skull of the large Tyrannosaurus “Stan” at the Black Hills Institute, South Dakota, before it was
auctioned off in 2020.
PHOTO BY THE AUTHOR

How Many Tyrannosaurs Were There?

We know Tyrannosaurus rex only from the Midwest of North America.

Close relatives of the king of dinosaurs, such as Tarbosaurus, also lived in
Mongolia or Zhuchengtyrannus in China. However, the genus
Tyrannosaurus lived in a spatially limited area. Although T. rex is so well-
known, it lived for only about 2 million years. How many individuals were
there during this time? This question cannot be answered reliably.
Nevertheless, colleagues at the University of Berkeley in California have
shown with an amazing experiment how one can get a rough idea. They
found an approach to this question with the help of the Monte Carlo
simulation, a method from probability theory and mathematical statistics
(stochastics). The Monte Carlo simulation attempts to solve analytically
unsolvable problems numerically. It involves taking samples of a distribution
that are determined by random experiments. To find out the abundance of
tyrannosaurs, a large number of variables must first be determined: In what
time period did T. rex live? How large was the area where tyrannosaurs
occurred, and what was the population density? How many offspring did a
tyrannosaur produce, and how many of these survived to sexual maturity?
How old did these animals live on average, and how fast did they grow? Did
they have an active metabolism, or were they ectothermic? How heavy was
an adult animal? What trophic level did the tyrannosaur occupy, and was
there perhaps a change in trophic levels at different stages of the animal’s
growth? How many generations of tyrannosaurs existed, and what is the
absolute number of skeletons that have appeared in the fossil record to date?
Then all this data was compared with data from present-day ecosystems, and
the results are mind-boggling. The Berkeley colleagues chose
Tyrannosaurus, of all dinosaurs, for this experiment because it is the best-
studied dinosaur, the time interval of its existence was short and the number
of individuals small since it was at the top of the food chain. Compared to
small carnivores or herbivores, the top predators in an ecosystem are very
rare. But what do the numbers say?
Tyrannosaurus roamed the Earth for about 1.2 to 3.6 million years before
the asteroid hit. The mean value is 2.4 million years. The Berkeley team
calculated the duration of a generation based on the proportion of sexually
mature individuals, their average number of offspring, their survival rate,
and their maximum lifespan. They arrived at a time span of about nineteen
years and thus found that about 127,000 generations of tyrannosaurs must
have existed.
The study puts the area where the tyrannosaurs lived at 888,000 square
miles. They estimate the average body weight of a sexually mature animal
was roughly 11,500 pounds. That’s about twenty-seven times the average
weight of a lion and about twenty times that of a tiger. Because
Tyrannosaurus was so heavy and had, like modern mammals, an active
metabolism, its energy requirements were quite a bit higher, which is why
there were far fewer individuals in an area relative to modern mammalian
predators. This is due to the fact that vegetation can only support a certain
number of herbivores, and the number of predators depends on the
abundance of available prey. Since an adult Tyrannosaurus cannot hunt
small animals and relies on large herbivores, this limits its abundance. The
researchers estimate that the population density of tyrannosaurs was only 16
percent of that of tigers and only 7 percent of that of lions. So, on a safari at
the end of the Cretaceous period, you would have been much less likely to
see an adult tyrannosaur than you would be to see a tiger or a lion today. It
would have been difficult to get a good snapshot of one. But you probably
would have encountered many more babies and juvenile animals—though if
there’s one thing we’ve learned from the Jurassic Park movies, it’s that you
wouldn’t want to run into a dinosaur in the wild anyway. In absolute
numbers, the Berkeley colleagues estimate that the population density of T.
rex ranged from 0.00058 to 0.14 sexually mature individuals per square
kilometer. With a mean of about 0.0091 individuals per square kilometer,
there were thus about twenty thousand to twenty-one thousand adults living
in the total area at any one time. Calculating the numbers, we find that a total
of about 2.66 billion adult tyrannosaurs lived. If one includes hatchlings and
young animals, it was certainly ten times more. But if you are familiar with
statistics and do the math yourself, you will find that the margin of error is
very high. If we follow the same calculation example, but assume the
smallest value (1.2 million years) for the reign of the tyrannosaurs and only a
value of 0.00058 for the frequency of individuals per square kilometer, then
there were only about 84 million adult animals over the entire period of their
existence. If instead we assume 3.6 million years and a frequency of
individuals of 0.14 per square kilometer, then there were an incredible 61
billion adult tyrannosaurs. Yet these are not the only variables that yield
imprecision. If we change, for example, the area size, the duration of a
generation, the average age of sexual maturity, the number of offspring, or
the average weight, we obtain completely different values.

Why Tyrannosaurus Was Not a Scavenger

Every once in a while, some scientist or another will argue that
Tyrannosaurus may have been a scavenger. A common argument in this
context is that its tiny forearms were useless for hunting prey. However,
paleontologists Kenneth Carpenter and Matt Smith clearly rejected this
claim through a detailed scientific description of Tyrannosaurus’s forearms.
Since more and more evidence has been found of this animal’s very active
metabolism, its rapid growth rate, and its incredible bite force, the voices
have quieted, although the scavenger argument still comes up occasionally.
But even in this case, mathematics can help. Indeed, it is absolutely
impossible to feed a six-ton tyrannosaur on carrion alone, as a dear colleague
of mine, paleontologist Chris Carbone, impressively demonstrated a few
years ago. I learned about his computational model in 2010 at a working
group meeting in Flensburg, Germany. A year later his article on the subject
was published. The calculations are not quite simple, as they require a lot of
research beforehand.
First, you have to estimate approximately how many tyrannosaurs there
were in an ecosystem, how much prey and how much carrion was in it, and
how long the carrion usually remained accessible as food. Then you have to
consider how large the area was where predators and prey coexisted.
Knowing these variables, the conclusions are clear. In this computational
model, too, we must use values based on observations in nature today. And
again, we seek an approach to an intractable problem by proceeding
analytically and solving the problem numerically. Weighing the pros and
cons of scavenging versus hunting is very important if we are to understand
the complex relationships in an ecosystem. This is especially true for the
interactions between carnivores of different trophic levels.
As one of the largest land-dwelling carnivores ever known,
Tyrannosaurus is a particularly popular subject of research, and in the more
than one hundred years since its discovery, there has been eager debate about
its diet. Was Tyrannosaurus an active predator or an obligate scavenger—
that is, an animal that was mandatorily dependent on carrion for survival? To
find an answer to this question, one must first understand that almost all
carnivores do not only actively hunt, but also eat carrion when given the
opportunity. Who would turn down a free meal that can be obtained without
any effort? Sometimes even herbivores ingest carrion to obtain important
minerals when the vegetation in their habitat is particularly low in nutrients.
 Nevertheless, few animals rely on carrion as a major component of their
diet because the occurrence of carcasses is unpredictable and no strategy can
be found for finding carrion at short intervals, regularly, and in sufficient
quantity. In all contemporary ecosystems, large carnivorous mammals hunt
primarily large vertebrates, and so it is more than likely that a giant
carnivore such as T. rex also hunted because, with its enormous weight and
massive energy requirements, it simply would not have been able to compete
with smaller predators as an obligate scavenger.
To illustrate this, we need to look at the environmental conditions of the
Late Cretaceous of North America and the size distribution of co-occurring
herbivorous dinosaurs, as well as the abundance of competing small
carnivores. In the fauna of the Late Upper Cretaceous of North America,
surprisingly, we find almost no medium-sized carnivores. There were either
small nimble predators with a body mass of less than fifty kilograms or huge
tyrannosaurs weighing up to eight tons. If we look at herbivores, the
distribution was much more balanced. There were also very large and heavy
representatives such as the horned dinosaur Triceratops, which weighed up
to twelve tons, or the duckbill dinosaur Edmontosaurus, with a body mass of
up to nine tons. At the same time, however, there were also much smaller
animals.
Triceratops and Edmontosaurus are the most common dinosaurs found in
the Hell Creek and Lance formations. Club-tail dinosaurs like the well-
known Ankylosaurus are much rarer there; they could weigh up to eight tons.
Most tyrannosaurs were found in the Hell Creek and Lance formations of
Montana, Wyoming, and South Dakota. But in recent years, remains of these
giant animals have also been discovered in New Mexico and Utah. Other
herbivores lived there—the largest among them and one of the last long-
necked dinosaurs of North America was Alamosaurus. Mass estimates of
these gigantic dinosaurs vary greatly. A 2020 publication cited about thirty-
eight metric tons. Besides these giants, there were also many smaller
herbivores, and most animals of this time were nestlings and juveniles
anyway. They were much more common than adults because dinosaurs laid a
lot of eggs, and their mortality rate was very high. Therefore, my colleagues
assumed that almost 50 percent of the herbivores weighed only between
fifty-five and eighty-five kilograms.
As mentioned earlier, only about twenty-one thousand tyrannosaurs
occurred at any given time over a vast area of 888,000 square miles. Based
on the expected frequency of carrion occurrence and the fact that carcasses
would have been quickly tracked down and consumed by smaller theropods
even before a Tyrannosaurus found it, a primary diet of carrion seems very
unlikely. Larger carcasses would have been very rare and highly contested,
making them an unreliable food source. My colleagues estimated the
potential carcass-discovery rate of smaller theropods to be fourteen to sixty
times that of an adult T. rex. These results suggested that T. rex was unable to
survive as an obligate scavenger and compete with small predators. Thus,
the prey it hunted was primarily large vertebrates, similar to many large
carnivorous mammals in modern ecosystems. But what numbers and
considerations go into this computational model?
First, we need to estimate how much carrion there was in the first place.
Today, a productive modern ecosystem like the Serengeti, in Africa,
produces twenty-five pounds of carcasses per square mile per day. If we
assume that carcasses were similarly abundant in Mesozoic terrestrial
ecosystems, there would have been a substantial number of dinosaur
carcasses in North America during the Late Cretaceous—enough to support
a single adult T. rex. However, if we estimate the distribution of carcasses of
different sizes in terms of the frequency they occur, it quickly becomes clear
that finding these carcasses would certainly have been a problem. Under the
simplifying assumption that all size classes of carcasses occur with the same
frequency as the corresponding live animals, we can easily calculate an
expected distribution. For this, we draw on the Serengeti production rate and
divide that rate among the size classes of herbivorous dinosaurs. Then we
calculate how many animals are in each class. If each carcass existed for
seven days before ultimately being consumed, most carcasses would be very
widely scattered indeed: on average, there would be only a single seventy-
five-kilogram carcass within a seventeen-square-kilometer radius, only a
single seven-hundred-kilogram carcass within one hundred and sixty square
kilometers, and only a single five-ton carcass within one thousand square
kilometers. Carrion from a twenty-five-ton herbivore would have occurred
on average only once in a five-thousand-square-kilometer radius. Thus, it is
clear that these carcasses were difficult, if not impossible, for individual
scavengers to locate without traveling very great distances. One animal
alone would never have managed to cover such distances and, more
crucially, the much more common smaller predators would have long
discovered and consumed the carcass in the meantime. Although Chris and
his colleagues base their calculations on a much lower T. rex population—
the authors calculate only about one third of the previously mentioned
twenty-one thousand tyrannosaurs (i.e., about sixty-six hundred to seven
thousand animals) in the total area—even with higher numbers the chances
of finding fresh carrion would have been almost hopeless.
One reason I talk about this, now quite old, article is that the variables
have recently regained relevance in the context of why there were no
medium-sized hunters in Tyrannosaurus-dominated ecosystems. From
today’s perspective, this unnatural frequency distribution of very large and
very small carnivores in an ecosystem where medium-sized predators are
completely absent was already noticed by paleontologists early on. For a
long time, scientists have tried to understand this phenomenon, but they have
not had a good explanation. We cannot see this evolution in ecosystems from
the Jurassic. On the contrary, in some of the most famous sites for Jurassic
dinosaurs, such as the Morrison Formation in the Midwest of the United
States, which we have already learned about, and the Lourinhã Formation in
Portugal, carnivores are distributed among all size groups, and lightweight
and heavy animals are similarly abundant. This is still true for some Lower
Cretaceous ecosystems, such as those of the Wessex Formation in England
and the Sao Khua Formation in Thailand.
Particularly fascinating in this context is the Huincul Formation from the
Upper Cretaceous of Argentina. There, besides huge carcharodontosaurs
that, like Tyrannosaurus, weighed more than five tons, middleweight classes
are also represented. Only in the Late Cretaceous, as soon as tyrannosaurs
became the dominant species, there was this very clear separation between
small predators and very large tyrannosaurs, and medium-sized predators no
longer occurred. Therein may lie the secret as to why the large dinosaurs
were unable to recover from the impact at the end of the Cretaceous.

Different Prey for Father and Son

In a 2011 article, paleontologist Chris Carbone compared different

ecosystems of the Late Upper Cretaceous of North America. Various
predatory dinosaurs lived in these areas. In his study, it is immediately
apparent that on the one hand there were very small predators, for example
the dromeosaurs with a body weight of thirty-five pounds, to the one-
hundred-and-forty-pound oviraptors, and then there was a huge size gap so
that the next largest carnivores weighed around twenty-four hundred pounds.
This was more than seventeen times the body weight of the smaller animals.
After these predators, there was another huge gap, as the adult tyrannosaurs
weighed an average of close to twelve thousand pounds, almost six times as
much. We don’t know any ecosystems today with such a size distribution.
But the authors of the 2011 article did not know that the twenty-four-
hundred-pound predatory dinosaur (it was Nanotyrannus) was not a separate
genus at all, but merely a juvenile tyrannosaur! This means that the actual
jump in mass ranges from one hundred and forty to twelve thousand pounds
—an almost eighty-five-fold difference! This makes this uneven distribution
all the more astounding.
People initially tried to attribute this phenomenon to the incompleteness of
the fossil record, and long assumed that medium-sized carnivores simply had
not yet been found. The argument has always been a bit specious, but at least
plausible, since carnivores are much rarer than herbivores in most
ecosystems, and therefore there were certainly fewer predators to be found
overall. However, since we have known many of the tyrannosaur sites for
one hundred and fifty years, it is clear that this cannot be the reason for the
lack of medium-sized species. Moreover, we see the same pattern in dozens
of sites in North America and Asia. So what is the reason? The secret lies in
the completely different biologies of tyrannosaurs and modern mammalian
predators. At the Kruger National Park in South Africa, there are more than
fifteen different species of predators. Lions, cheetahs, spotted hyenas and
brown hyenas, leopards, African wild dogs and African wild cats, jackals,
caracals, servals, among others, can be found there. Moreover, there are
African civets, bat-eared foxes, Cape foxes and black-footed cats. And their
body masses range from 330 to 550 pounds for lions to only 2.9 to 4.2
pounds for black-footed cats. In between, all weight classes are present:
cheetahs and leopards weigh between 46 and 159 pounds; hyenas between
84 and 141 pounds; caracals, jackals, and servals between 14 and 42 pounds;
and the small cats and foxes between 2.9 and 14 pounds. Many weight
classes are thereby jointly occupied by several species together.
In the dinosaur ecosystems in the Jurassic and up to the early Lower
Cretaceous, we still see a greater diversity of species in predatory dinosaurs
over fifty kilograms. So medium-sized predatory dinosaurs were also
represented there and thus all weight classes were filled. From about 83
million years before our time this changed in East and Central Asia and in
western North America (Laramidia). There, tyrannosaurs occupied a
monopoly position, and medium-sized adult predatory dinosaurs with a body
mass between one hundred and ten and eleven hundred pounds were very
rare or even absent entirely, even if they had still been present in these
regions before the Campanian. Although predator diversity declined, there
was no decrease of prey diversity. The missing food niches in theropod
assemblages were occupied by juvenile and subadult tyrannosaur species
that were functionally distinct from their adult parents. Herein lies the
fundamental difference between the large carnivorous mammals of today
and Tyrannosaurus: because mammalian young are suckled by their
mothers, they indirectly consume the same food as their mothers. Babies of
big cats are relatively large and grow quickly. In addition, the cubs are
protected by the pride and do not have to hunt for themselves until they are
big and strong enough to do so. So, they eat the same food as their parents,
even though they are still too small to hunt this prey under their own power.
A lion thus always occupies the same ecological niche from birth and is also
always positioned on the same trophic level. With tyrannosaurs it was
different. Because dinosaurs hatched from eggs that could not grow to any
arbitrary size, tyrannosaur hatchlings were tiny compared to their parents. A
tyrannosaur egg is about sixteen to eighteen inches tall and a hatchling is
about twenty-seven to thirty-two inches long. A baby T. rex weighed about as
much as a small-to medium-sized dog. The mother of these babies was up to
one thousand times heavier. We now know that tyrannosaurs had an active
metabolism and grew very quickly. However, because the difference in
weight between a hatchling and an adult dinosaur was so great, it still took
almost eighteen years for a T. rex to become sexually mature. Based on this
long time span, we must assume that tyrannosaurs occupied various feeding
niches during their lives, some of which were otherwise occupied by
medium-sized theropods prior to that time. Tyrannosaur juveniles occupied
these niches for themselves during their long growth phase because their
body proportions changed considerably, and they had to develop new
hunting strategies and kill different prey during different life stages. This
also means that the novel physique of tyrannosaurs was superior to that of
other medium-sized predators; they could not otherwise have outcompeted
their rivals. Even a juvenile Tyrannosaurus, which had about 10 percent the
biting force of an adult animal, was nevertheless stronger and faster than
other predatory species.
It thus ensured the evolutionary success of his species and gradually
replaced all other carnivores. Some raptors still existed, for example, in the
Hell Creek Formation, but these were extremely rare and do not play a role
in the consideration of faunal assemblages as a whole. One of these was
Dakotaraptor, which weighed between 220 and 1,100 pounds. It was about
the same size as the raptors in the Jurassic Park movies and looked similar.
In the sixth installment of the film series, it is even introduced as a separate
species, distinct from the other raptors. In the Hell Creek Formation, this
predatory dinosaur may have occupied a special ecological niche not
occupied by the young tyrannosaurs, and its pack hunting behavior probably
made coexistence with much larger tyrannosaurs possible. It may have killed
prey that was still too large for juvenile T. rex. Besides it, there were some
troodontids that weighed between 110 and 220 pounds. Whether these
actually competed with tyrannosaurs for resources is questionable, since they
were most likely omnivores and did not fulfill much of their dietary needs
with meat. Overall, coexistence in an ecosystem is only stable if the prey
selection of different genera is different. Thus, we see a decline in species
richness among predatory dinosaurs at the end of the Cretaceous, but not in
the number of ecological niches. This has to do not only with the extreme
increase in size of the apex predators, but also with the major changes in
their physique, which were so extreme that experts speak of a “secondary
metamorphosis” of the tyrannosaurs. They changed their appearance so
much that the young animals “functioned” differently than the adults.
Although both belonged to the same species, they occupied functionally
different niches. This is also a reason why juveniles of Tyrannosaurus were
not even recognized as such in the beginning but were described as a
separate species. Not only did their whole body structure change, but also
their proportions. The initially rather elongated skull, which looked like that
of Allosaurus, became increasingly higher in vertical extension, which
provided more space for larger and thicker jaw muscles and, throughout
ontogeny, ensured that the bite force increased more and more. The ratio of
arm length to leg length also changed; in juveniles, the arms were still
relatively long but eventually stopped growing while everything else
continued to enlarge. This means that a juvenile may have used its arms to
catch, injure, or kill prey because its jaws were not yet strong enough to do
so. What’s more, at some stages of development, the animal’s body and tail
grew faster than its legs, which is why the adult dinosaurs look beefier and
more massive, while the juveniles look slimmer and nimbler. So young
animals had proportionally longer arms; their humerus was about 40 percent
the length of the femur, whereas that of adults was only 30 percent. Once
their jaws became stronger, however, they no longer needed to hold or injure
their prey with their hands. Thus, a species occupies different food niches
throughout its life. This is called “niche assimilation,” and we can observe it
in all large dinosaurs.
We observe something similar in the long-necked and duckbill dinosaurs.
The Edmontosaurs from the Hell Creek Formation underwent a striking
increase in size during ontogeny. Their body structure was functionally
different at different stages of their lives. They inhabited a wide variety of
habitats during their ontogeny and occupied as many food niches as the
smallest grazing artiodactyl, the dwarf antelope, to the largest herbivore, the
African elephant. This represents the entire range of ungulates and
proboscideans of the Serengeti.
We find no analogy to niche assimilation in tyrannosaurids in the animal
kingdom today. Compared to the carnivorous mammals of Kruger National
Park, it would be as if a tyrannosaur, in the course of its life, grew from a
small black-backed jackal to an animal fifty times heavier than an adult lion!
These niche shifts happened abruptly in tyrannosaurids. With increasing
size, transitions from one phase to the next were rapid, and transformations
in the skeleton and body structure were profound, which in turn changed the
way prey were acquired relatively abruptly, too.
In summary, large theropods, unlike carnivorous mammals, occupied
multiple niches throughout their lives. This resulted in taxonomic
impoverishment, but not a reduction in niche diversity, because juvenile
large predators functioned as independent ecological morphospecies. This
explains the absence of medium-sized predators in the uppermost
Cretaceous. Their niches were filled by the adolescent mega-predators. We
can observe this gap in size distribution on different continents, in different
habitats, and in different species when the apex predator is from the
tyrannosaurid clade. This clearly indicates that this phenomenon is not an
artificial effect caused by gaps in the fossil record or a collection bias, but
reflects competition for resources in habitats where most niches were filled
with juvenile tyrannosaurids. This specialized structure of dinosaurs is
related to their extreme size and to the fact that they were egg-laying. Their
relatively small eggs hatched into small young that grew very rapidly,
resulting in this ontogenetic niche shift. Despite the high mortality rate, the
high number of offspring ensured a large juvenile population that
morphologically and functionally replaced meso-carnivores. Juveniles were
fast and agile and could run with endurance; their parents, on the other hand,
had a much higher bite force. This niche separation was necessary because
small prey would not have been sufficient for the large tyrannosaurs to meet
their energy requirements. In return, the young tyrannosaurs lacked the
strength to kill large prey such as adult duckbill or horned dinosaurs, on
which the adult tyrannosaurs fed.

I Am the King and Not a Peacock: Did T. rex Have Feathers?

When the first predatory dinosaurs with feathers were discovered in China
about twenty years ago, it was a sensation. Over time, more and more finds
came to light that showed evidence of plumage in theropods and triggered
speculation about whether T. rex was covered in feathers also. Since we have
now found fossilized integument from tyrannosaurs, we can answer this
question.

In biology, the term “integument” refers to the entirety of the

dermal layers of animals and humans, including the hair,
feathers, spines, scales, and bony plates formed in the skin.

Fossilized remains of T. rex and other tyrannosaurids (Albertosaurus,

Daspletosaurus, Gorgosaurus, and Tarbosaurus) show that large and heavy
tyrannosaurids did not have feathers, but instead had a scaly, reptilian-like
skin. At least this is true for all adult tyrannosaurids, on the majority of their
bodies. We cannot completely rule out the possibility that these dinosaurs
wore individual decorative feathers as ornaments. However, there are other
Tyrannosaurus relatives that were indeed completely covered in feathers,
such as Yutyrannus from China. Its name is a mixture of Mandarin and
Latinized Greek and translates to “feathered tyrant.” It is the largest
feathered predatory dinosaur disovered to date. Three almost completely
preserved specimens of this animal have been found so far. Of these, one is
fully adult, one is a subadult, and the third is a juvenile. Although its name
suggests it might be, Yutyrannus is not a direct relative of T. rex, but evolved
in a sister group. The fossils of Yutyrannus are about 130 million years old
and show that gigantism evolved independently (i.e., convergently) at least
twice within these sister groups. Here there is a superfamily called
Tyrannosauroidea. The higher order is the Coelurosauria, the so-called
hollow-tailed lizards. The superfamily Tyrannosauroidea includes two
families: the Proceratosauridae, to which the Yutyrannus belongs, and the
Tyrannosauridae that includes the Tyrannosaurus rex. The latter are
commonly referred to as tyrannosaurids, and they include all the other
known tyrannosaurs mentioned above (Tyrannosaurus, Albertosaurus,
Daspletosaurus, Gorgosaurus, and Tarbosaurus). There is currently no
evidence that adult Tyrannosauridae had any plumage. They are uniformly
covered with horny scales on the neck, abdomen, hips, and tail. If these
animals had feathers, they most likely occurred on the dorsal part. We
currently believe that tyrannosaurids lost their abundant pennaceous feathers
in the late Lower Cretaceous (the Albian) about 110 to 100 million years
ago. Some researchers initially assumed that this loss had to do with the
warmer paleoclimate in which the tyrannosaurids lived. However, this is
now considered very unlikely. The geographic range of the tyrannosaurids
was vast, which is why their representatives lived in extremely diverse
climatic conditions. A special example is Nanuqsaurus, “the polar bear
lizard,” because its remains were discovered in the Prince Creek Formation
in the extreme north of Alaska. It must have been very cold there during the
winter months, and yet there is still no evidence that this animal possessed
feathers. Thus, the loss of feathers is apparently not related to the
paleoclimate.
According to the first description, Yutyrannus was about thirty feet long
and weighed roughly three thousand pounds. Other authors assume that the
adult animal was only about twenty-five feet long and weighed twenty-four
hundred pounds. This may not seem like much of a difference at first glance,
but perhaps that is why this animal still had feathers and the larger
tyrannosaurids did not. Albertosaurus, Gorgosaurus, and Daspletosaurus
weighed about two and a half tons each (some estimates even go up to four
tons for the heaviest specimens). Tarbosaurus is thought to have weighed as
much as about four tons, and T. rex averaged perhaps five to six tons, with
the very largest animals possibly weighing more than nine tons.
 When certain groups of animals undergo an evolution toward gigantism,
the ratio of body surface area to body mass changes dramatically. This is
because the amount of surface area increases to the second power, but the
volume increases to the third power. Such large and heavy animals were
indirectly gigantothermic—that is, their bodies maintained a constant body
temperature regardless of the outside temperature, because cooling their
body through conduction would have taken far too long. Now, if a thick
insulating layer had been added, such as a dense plumage, there would have
been hardly any heat loss through conduction. Conversely, this would have
meant that an animal with a large body mass could quickly overheat. In
sauropods, this problem was solved by a complex system of air sacs, and
mammals have sweat glands to regulate body temperature. In the case of the
heavy tyrannosaurids, the development to gigantism in the course of their
evolution had to have inevitably led to the loss of their plumage, so that they
did not overheat.
Yutyrannus, which was still fully pennate, apparently had not yet exceeded
the threshold at which a full coat would have caused overheating, with its
body weight of about one ton. There may have been other reasons that
Yutyrannus had feathers as opposed to Tyrannosaurus—Yutyrannus lived in
a more densely forested habitat than Tyrannosaurus and thus may have
needed more protection and thermal insulation—but the deciding factor was
the risk of overheating, to which Tyrannosaurus was more exposed. In
addition, tyrannosaurids had longer legs than their close relatives, suggesting
a lifestyle still seen in ratites today. They cover long distances in open terrain
and can sometimes run very fast. This behavior is extremely energy
consuming and may have increased the risk of overheating during prolonged
chases.
However, because tyrannosaurs are descended from feathered ancestors
and almost all hollow-tailed lizards (Coelurosauria) had pennaceous
feathers, we now assume that the horny scales of T. rex are not comparable
to the scales of crocodiles (i.e., not homologous), but that they evolved
secondarily from feathers. If anyone had claimed that thirty years ago, they
would have been considered simply crazy. But the same is true for the scales
on the legs of some modern birds: they also evolved from feathers. This
secondary transformation of leg feathers into scales is, in the case of today’s
birds, an adaptation to wading in water—for example, in storks or herons—
or to crouching and balancing on thin branches.
 So adult tyrannosaurs were scaly. But what about chicks? Were they
feathered or did they already have scales? The fact that adult animals were
covered with horny scales does not necessarily rule out the possibility that
the young still had feathers, but that these were lost in the course of their
ontogeny. However, such a change from feathers to horny scales is
unprecedented in the animal kingdom today, and therefore it will remain
ambiguous until we find the first remains of a baby Tyrannosaurus with
integument! I would not be surprised if a loss of feathers coincided with a
dramatic growth spurt, during a period when the young were transitioning
from one ecological niche to another. The small hatchlings may have
benefited from the protection of the dense forest and found their prey there,
for which they needed feathers, while the juvenile tyrannosaurs went hunting
in open terrain and therefore no longer needed them. This scenario is
plausible because the juvenile tyrannosaurs, relative to their bodies, had
longer legs than the adults and were fast runners.

Short Arms Are Normal but Long Ones Are Strange

Many jokes about T. rex allude to its short arms. They were downright tiny
compared to its otherwise enormous proportions. Initially, only a humerus
and no forearm or digits were found from T. rex. Henry Fairfield Osborn, at
the first public exhibition of the skeletal reconstruction in 1915, had
mounted three-fingered hands on it, as had been seen in Allosaurus. But he
should have known better, because the closely related Gorgosaurus was
already known at that time, and it had two fingers on each hand. And
although there was later no doubt that T. rex also had only two fingers on
each hand, it was only a specimen discovered in 1989, which is now on
display in Montana, that provided unequivocal evidence of this, because in
this fossil the entire arms had been preserved for the first time. Today it is
clear that all tyrannosaurids were two-fingered.
But why were the arms of T. rex so short compared to its body size? They
measured just around three feet, and some of my colleagues describe them as
rudimentary. But we shouldn’t make fun of it—muscle attachment scars on
the humerus, in fact, show that Tyrannosaurus’s biceps were very strongly
developed. The bones of their arms had extremely thick cortical bone,
showing that the animal could withstand heavy loads. The biceps of an adult
T. rex were so strong that it could lift four hundred and forty pounds with
them. Other muscles in the upper arm, such as the brachialis muscle, worked
in concert with the biceps to strengthen elbow flexion. So, T. rex would not
only have won against any human in arm wrestling—its muscle strength
would have been enough to simply rip out a human’s entire arm.
While human upper arms can rotate three hundred and sixty degrees at the
ball-and-socket joint of the shoulder, tyrannosaurs had very limited range of
movement. They could only pivot their arms forty degrees at the shoulder
and forty-five degrees at the elbow. Biomechanical analyses of the massive
arm bones, extreme muscle strength, and limited range of motion suggest
that the short arms evolved so that these dinosaurs could hold on to their
prey even in the face of massive opposition. The large muscle attachments
on the humerus caught Osborn’s attention as early as 1906 and he correctly
concluded that these arms must have been very strong. He assumed that they
were used to hold a partner during mating. In the 1970s, it was assumed that
the arms were important when standing up from a prone position to prevent
the animal from sliding forward while pushing with its hind legs. All three
considerations are plausible and need not be mutually exclusive. In 2021,
however, Professor Kevin Padian of Berkeley argued that the reduction of
arms in tyrannosaurids had no special function but was rather a secondary
adaptation. According to him, the arms became smaller as the animals’ skulls
became larger and their jaws stronger. This served to prevent bites and
serious injuries when, for example, a pack of tyrannosaurs fought over the
same prey. But whether this is tenable remains questionable. In Albertosaurs
we see such pack behavior. They lived at least partly together with
conspecifics of the same age, but for Tyrannosaurus so far there is no
evidence for such behavior.
The idea that the arms were used as weapons during hunting and the claws
were used to slash prey, however, seems more than questionable to me. The
arms are much too short for that and their range of motion is, as mentioned
above, much too limited. The claws on their fingers could have inflicted long
and deep cuts on the prey, but Tyrannosaurus probably could not have
targeted vital or vulnerable areas with them.
But if you think that the tyrannosaurs had the smallest forearms relative to
the rest of their bodies in the dinosaur kingdom, you are wrong. The arms of
Carnotaurus and Abelisaurus from Argentina or those of Majungasaurus
from Madagascar are much shorter. The three carnivores belong to a group
that we call abelisaurians and are found mainly in the southern hemisphere.
 This probably seems strange to us because we don’t know any other
dinosaurs today except for birds. The extremely short arms seem strange to
us, even if, conversely, it is actually the anatomy of modern birds that is
special. With them a clear decrease of body size took place with a
simultaneous increase of relative arm length. The wings are unusually long
for dinosaurs. The evolution of birds toward a small body size and long arms
probably made active bird flight possible. Indeed, there is conversely an
evolutionary trend of negative forearm allometry in theropods that are not
yet birds, where larger species often have relatively short forearms. This
contradiction can be explained by the fact that the forearm bones, i.e., the
ulna and radius, were disproportionately long early in the body evolution of
predatory dinosaurs. Thus, the longer a juvenile lived, the longer its humerus
and femur became relative to the forearm, which was accompanied by a
significant change in arm function. Because tyrannosaurs were not fully
grown until they were about eighteen years old, but their arms stopped
growing much earlier, the arms of adult animals appear very short. This was
not yet noticeable in T. rex teenagers, because a three- or four-year-old T. rex
already had arms as long as its parents, and they simply stopped growing
after that. A large abelisaur or T. rex killed its prey with its mouth and did not
need strong arms. Consequently, the transition from one food niche to the
next also determined arm development. Conversely, small theropods needed
arms to catch prey and, in the case of birds, to be able to fly. The negative
allometry of the arms is thus a signal for evolutionary developments, which
are also reflected in ontogenetic trends. In modern birds, the growth of arms
is related to changes in movement and behavior during ontogeny—that is,
bird flight. The proportionally longer arms of birds are a juvenile trait that
adult birds retain. We have also seen the same phenomenon in the dwarfing
of Europasaurus, supporting the notion that the evolution of modern birds
was driven by paedomorphosis. Paedomorphosis can be achieved by
accelerating sexual maturity relative to the rest of development, or by
delaying physical development relative to reproductive capacity. That’s why
birds basically look like baby dinosaurs.

Boy or Girl: Was Sue Female?

The T. rex at the Field Museum is named Sue—but is the animal really
female? It’s hard to tell. Distinct sexual characteristics in dinosaurs are
usually lost during fossilization. But one particular bone tissue is found only
in female birds, and in all other female dinosaurs—medullary bone tissue. It
is a special, derived tissue that lines the internal medullary cavity of females
just before they lay eggs. Medullary bones in extinct dinosaurs suggest
similar reproductive strategies and serve as an objective method for their sex
differentiation. However, there is a catch: it only helps us identify females
that were sexually mature and about to lay eggs at the time of their death.
This is because the medullary bone served as a calcium reservoir, which the
females needed for eggshell formation. In Sue, we do not find this bone
tissue, which does not necessarily mean that she was a male. It may simply
mean she hadn’t laid eggs shortly before her death. Medullary bone tissue is
known from two other tyrannosaurs, however. One of these animals is on
display at the Museum of the Rockies in Bozeman, Montana. It is nicknamed
B. rex and at the time of its death was about eighteen years old and already
weighed three tons. Of B. rex less than 40 percent of the skeleton is
preserved, but its skull is nearly complete. The medullary bone tissue came
from the medullary cavity of its femur, from which other spectacular
discoveries were made, too. In fact, contrary to popular belief, this
tyrannosaur showed that fossil bones can sometimes contain original cells,
blood vessels, and structural tissue still composed of its original proteins. In
this case, this was due to the extraordinary deposition and preservation of the
animal after its death. The carcass of B. rex decayed in a brackish estuarine
channel and was buried under sand in an oxygen-rich environment, rapidly
solidifying. As a result, no further chemical decay occurred. Much research
will certainly be done on this in the coming years.
The other female is housed at the Burpee Museum of Natural History in
Rockford, Illinois. It was about fifteen years old at the time of its death, still
a teenager. Fossils of juvenile tyrannosaurs are extremely rare, but the
Burpee Museum has not one but two of them, both found in the same region
of Montana. The more complete specimen of the two teenagers, nicknamed
Jane, was slightly smaller and, at about thirteen years old, a bit younger.
However, we do not yet know if Jane was a female or even sexually mature.
Paradoxically, medullary bone tissue was detected in the larger animal,
which bears the male nickname Petey. Jane, by the way, is that specimen that
was thought to be the new species Nanotyrannus lancensis. However, since
it turned out that it was actually just a young T. rex, the animal is called
Tyrannosaurus again.
 The King, the Queen, and the Emperor

With all the confusion about gender and dramatic changes in body structure,
it is not surprising that some of my colleagues believed that there may have
been several different species of Tyrannosaurus. The species Tyrannosaurus
rex is widely known, but researchers have long observed that some animals
were more robustly built and some had wider pelvises than others. Some
paleontologists interpreted these as gender-specific traits. They argued that
the narrow pelvis was indicative of males and the wider one was indicative
of females, because a wide pelvis would certainly be advantageous when
laying eggs. Others saw the high variability as a sign that there must have
been several different species. This hypothesis does not seem to me to be
completely far-fetched; after all, as we have already learned, Tyrannosaurus
existed on Earth for about 1.2 to 3.6 million years. This time span is
sufficient for it to have split into several species. A 2022 article states that
there was originally one species of Tyrannosaurus, which later gave rise to
two new ones. The authors noted that many footprints and skeletons of T. rex
showed considerable variability that had not yet been studied in detail
stratigraphically. This meant, they said, that it is not yet known whether the
differences might not be related to a different temporal occurrence of the
respective finds. In studies of more than three dozen specimens of T. rex, the
authors found evidence of a remarkable disproportionate variation in
tyrannosaurs, which they believed could not be explained by different
growth phases or sexual dimorphism alone, but by the fact that various
forms occurred in chronological succession and were found in different rock
strata of different ages. They cited the robustness of the skeleton and the
varying number of small incisor-like teeth in the lower jaw as the most
important arguments for their theory. Thus, at the beginning of the
tyrannosaur evolution, there was one robust species with two small teeth in
the front and much larger teeth farther back in the lower jaw. The robustness
of the animals was defined by the circumference of their femora. The
original form of T. rex is thought to have later split into two new species that
had only a single small tooth in front of the larger teeth in the lower jaw. One
of these new species still had robust femora and the other one had more
slender ones. The two new species then replaced the original species.
 When a new species develops from an old one, it is called
anagenesis. When an original species has split into two new
sister species occurring at the same time, a so-called
cladogenesis took place. Something like this can happen when
two populations are spatially separated and then adapt
differently to their respective habitats.

The first skeleton of a T. rex ever found was not at the same time the
geologically oldest. It was a more derived form as a result of the anagenesis,
and a robust one at that, but bearing only one small tooth in the anterior
mandible. However, according to the rules of nomenclature, the specimen
which was described first must retain its original name, so the authors gave
this species the name Tyrannosaurus rex. For the oldest robust form with
two small teeth at the tip of the lower jaw, they later gave the name
Tyrannosaurus imperator, while they renamed the more gracile new species
Tyrannosaurus regina. I like all of these names very much. Tyrannosaurus
rex, as we recall, means “king of the tyrant lizards,” Tyrannosaurus
imperator means “emperor of the tyrant lizards,” and Tyrannosaurus regina
means “queen of the tyrant lizards.” According to this new naming scheme,
Sue would be a Tyrannosaurus imperator, and a Tyrannosaurus regina can
be seen at the National Museum of Natural History in Washington, DC.
However, it is uncertain whether these names will catch on in the long run. I
found the distinction of these three species based on only two characteristics
very questionable, and shortly after the publication of the article in question,
many experienced colleagues shared my concerns. There are doubts about
the coherence of some arguments. If one postulates such a distinction of
different species, one should clearly prove it. Thereby the assumption of a
species succession over a period of 2.4 million years is quite plausible. In the
case of Triceratops, the species Triceratops horridus was followed by the
species Triceratops prorsus in the same period. On the other hand, there are
very many skeletons of Tyrannosaurus that have hundreds of distinct
skeletal features by which its species can be described. To derive three
species on the basis of only two features does not seem very convincing to
me. But the authors did not succeed in finding any more features among the
many that support their hypothesis. In fact, femur circumference varies
widely among tyrannosaurs, and other authors have shown that there are not
two size clusters with thick femora on one side and slender femora on the
other, but that there is a whole range of bones of different thickness, with
some occurring somewhere in the middle.
The tooth variation is a groundless argument. With mammals, this
characteristic would have perhaps been convincing, since they usually have
a strict tooth formula so all animals of the same kind always possess the
same number of teeth. Humans, for example—so long as they brush their
teeth thoroughly—always have two incisors, one canine, two premolars, and
two molars per quadrant. As they get older, they then grow one more molar
per quadrant, which we call wisdom teeth. Reptiles, on the other hand, have
very variable tooth formulas that allow for much more plasticity. Two
animals of the same species may have more or fewer teeth in their jaws. To
base different reptile species on different tooth formulas therefore does not
seem very reliable to me. That the distinguishing features are also inadequate
becomes clear from the example of the New York Tyrannosaurus, which is
one of the most complete specimens in the world. In this animal, the skull,
ribs, pelvis, and the entire spine up to the tip of the tail are almost
completely preserved. It is only missing its thighs. Consequently, it would be
impossible to assign this animal to a Tyrannosaurus species because we do
not know the circumference of the femora. It is not surprising that some
colleagues refuted the corresponding article only half a year after its
appearance. Perhaps someday we will actually be able to determine
differences that will make it necessary to divide the genus Tyrannosaurus
into several species. But so far this is not necessary, and so the name of our
favorite dinosaur remains with us for the time being: Tyrannosaurus rex!

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 Chapter 11

Movement Captured in the Stone:

What Footprints Tell Us

The German Dinosaur Highway

Much of what we know about dinosaurs, we have learned by closely
examining their bones. The size of the bones suggests the overall size of the
animals, and the circumferences of their arm and leg bones give us a rough
idea of how heavy dinosaurs were. Processes and muscle attachment scars
on the bones tell us how strong the muscles were and where they ran across
the body. The internal bone structure tells us if the animals had an active
metabolism, how fast they grew, and when they were sexually mature. Bone
cells allow us to draw conclusions about genome size. Medullary bone
helps us determine their gender. From the feathers of enantiornithines found
in amber, from the dermal bones of the armored dinosaur Borealopelta
found in the oil sands of Canada, and from the scales of dinosaur mummies
and tyrannosaurs, we get an accurate picture of the dinosaurs’ appearance
and their biology. As a young boy, I never dreamed that one day we would
find out what color dinosaur eggs were and what the cloaca or lungs of
these animals looked like. Our picture of the prehistoric lizards is becoming
clearer, and it’s one of dynamic, agile, and intelligent animals. Today, we
know that some dinosaurs displayed brood care behavior and were caring
parents. The dinosaurs of my childhood were clumsy, lumbering, and all
gray. Today, we see them on Apple TV in the most dazzling rainbow colors
and complex food webs. But even though the fossilized bones, scales, and
feathers reveal a lot about the life of the dinosaurs, they tell us little about
the everyday lives of these animals. They cannot capture a moment in the
lives of dinosaurs. Only the so-called trace fossils can do that. They are like
a snapshot frozen in time over millions of years. Unlike bones, which are
often transported, trace fossils remain exactly where they were produced.
Scientists call such fossils, which remain in place, “autochthonous.” Other
examples of autochthonous fossils are fossilized reefs. If we find dinosaur
tracks, we know that at some point in Earth’s past, a living animal walked
in that exact spot. We can tell whether the animal moved fast or slow, was
heavy or light-footed, whether the ground was sandy or muddy, and
whether the animal walked alone or as part of a herd. Because sauropod
tracks are autochthonous, they provide unambiguous information about the
actual habitat of the track maker. The giant tracks of gigantic sauropods
occur in sediments from the Late Triassic to Cretaceous periods around the
world. In the fossil record, they are the second most common evidence for
the long-necked giants. Tracks are important because they reveal
anatomical details and locomotion patterns of the track maker that are
otherwise difficult to determine.
Some of the best-known tracks are in mudflat deposits of the Paluxy
River near Glen Rose in Dinosaur Valley State Park in Texas. More
dinosaur tracks were discovered in the early 1980s, in the Kugitangtau
Mountains of eastern Turkmenistan at an elevation of five thousand feet—
sometimes called the “dinosaur plateau.” They are among the longest
continuous tracks ever found. New methods in studying the tracks are
helping us to better understand dinosaur biology. The principles of soil
mechanics inspired a field experiment with an African elephant whose
footprints were used to infer the animal’s weight. The geometry, depth, and
diameter of the footprints, in combination with soil mechanical analyses,
were used to calculate the mass as reliably as possible. Subsequently, it was
intended to apply this method to dinosaurs as well. The experiment was
conducted at the Wuppertal Zoo with a female elephant. The elephant was
weighed, and the track was prepared so that deformations in the sand could
be measured accurately. The geometry of the footprint was determined by
laser scanning and the velocity was determined with digital image
correlation. The properties of the soil were determined in advance by
experiments in the laboratory. Finally, for the reverse calculation of the
elephant’s weight, the finite element method was applied. The study showed
that footprint geometry, together with theoretical considerations of
subsurface loading, is well suited for predicting the weight of the track
maker. Because this method can be used to reliably measure the applied
additional load on certain substrates and calculate an animal’s weight with
an error rate of only about 15 percent, it will certainly play a major role in
dinosaur mass calculations in the future. But the process is complex
because several properties of the footprints and the substrate must be taken
into account before reliable results can be obtained for the fossils. In
addition, researchers must account for geological processes that may have
altered the original shape of the sediment. The method only works reliably
in sandstone because sand is not further compressed during its diagenesis to
sandstone, whereas in mudstone, measurement accuracy is compromised. A
grain size analysis and a detailed sedimentological investigation are
therefore important in advance.

An ideal place to test this method was the dinosaur track site of
Münchehagen, a natural monument in a former quarry, where hundreds of
sauropod and theropod tracks can be found over an 160,000-square-foot
area. There—about 140 million years ago, at the beginning of the Lower
Cretaceous—the dinosaurs left their footprints in the soft subsoil of an
estuarine delta. These tracks can be traced further in the adjacent quarry.

The Dinosaur Tracks of Lommiswil

Sometimes, tectonic movements of the Earth’s crust can cause dinosaur

tracks to be exposed vertically, as can be observed in Switzerland, halfway
between Basel and Bern, near Lommiswil. The excursion to Frick, where I
saw the plateosaurs, had also led me to the track site of Lommiswil. The
tracks there were only discovered about thirty years ago, although the
nearly vertical rock face has been exposed for eighty years. One would
simply not expect the sauropod tracks, which are showcased here, to be
found in marine deposits. The tracks are from the Kimmeridgian, which
corresponds to the “White Jurassic,” and are dated to 157.3 to 152.1 million
years before our time. They are thus only slightly older than the tracks from
Münchehagen. In the Upper Jurassic, sea level fluctuations apparently led
to temporary silting at this site, allowing dinosaurs to enter the otherwise
water-covered area. The partially visible mud cracks indicate a period of
drying of the water body. They form when the mud contracts as a result of
receding water. At that time, the sauropod dinosaurs walked through
calcareous mud plains, deposited in tidal pools, which are now known as
the “Solothurn Turtle Limestone.” Indeed, sea turtle fossil finds are also
known from there. The traces of the hind feet of these dinosaurs have a
diameter of more than three feet, and those of the front feet are somewhat
smaller, rather horseshoe-shaped, and less clearly visible. Thus, the front
legs did not sink as deeply into the mud, as the main weight of the animal’s
body rested on its hind legs. These animals were toe-walkers, and their toes
were embedded in a large pad of gristle similar to the shock-absorbing pad
of today’s elephants. The tracks of the hind feet of the sauropods have a
circular to oval shape and no claws are visible. We can therefore assume
that the trackers were probably not Diplodocus relatives, since three claws
on the hind feet are characteristic of this group. Depending on how soft the
mud that the sauropods walked on was, this detail might simply not have
been preserved, even though the animal may have had claws after all. A
better indication is the width of the tracks.
 The steep face with sauropod tracks in Lommiswil, Switzerland.
PHOTO BY THE AUTHOR

Although Diplodocus is a long-necked dinosaur with one of the longest

necks, it was quite delicately built. It had a narrow pelvis and was more like
a greyhound among sauropods rather than a bulldog. The narrow hips meant
that the legs were cantered under the body, and the animal put one foot right
in front of the other while alive. If a line were drawn running between its
left and right footprints, it would intersect both footprints. Later in the
evolution of long-necked dinosaurs, the titanosaurs, on the other hand, had
such massive bodies that they walked with their legs wide apart. The tracks
at Lommiswil show such a broad-gauge gait.
 There are broad-gauge trackways and narrow-gauge
trackways. These are terms derived from railroads, where
there are narrow-gauge and broad-gauge tracks.

Titanosaurs did not yet exist in the Upper Jurassic, but the brachiosaurs
also already walked with a broad-gauge gait. Their tracks were not caused
by a herd moving together in the same direction, but by individuals that
crisscrossed. Therefore, no clear predominant walking direction can be
recognized. The animals moved very slowly and were probably searching
for food. Calculations showed a walking speed of only two to three miles
per hour. Spore findings show that various conifers and fern species were
available as food sources for these animals. Conifers were well adapted to
the prevailing arid climate, but also to brackish conditions. In addition, the
sauropods may have fed on trees with water-storing leaves and herbs that
grew on the tidal flats. The tracks usually stand out from the rest of the rock
by their lighter coloration because the tracks were subsequently covered
and infilled with carbonate plugs. However, as the escarpment is exposed to
weathering, the tracks are beginning to fade and will become less and less
distinct in the future.

The Track Site of Cal Orck’o in Bolivia

Some of the most impressive dinosaur tracks are located near the city of
Sucre in Bolivia. At the Cal Orck’o mountain, a paleontological site, there
is a cliff with thousands of footprints. It is about two hundred and sixty feet
high, four thousand feet wide, and has an incline of seventy-three degrees.
The tracks come from hundreds of different turtles, crocodiles, small
lizards, but also from sauropods, theropods, and ankylosaurs from the
Upper Cretaceous period. The wall in central Bolivia has been known since
1968, and recent work shows that track sites also exist elsewhere in Bolivia,
occurring in several strata from the Campanian to the late Maastrichtian
(83.6 to 66 million years ago). They all belong to a mega-track complex
that extends from southern Peru across the central Bolivian Andes to Salta
Province in northern Argentina. This is probably the largest assemblage of
dinosaur footprints in the world. The main track area is about two hundred
and seventy thousand square feet and was mapped in 1998 using heavy
mountain-climbing equipment. The track-bearing strata show episodic soil
formation, stromatolites, and storm deposits. They formed in calcareous
lake deposits that repeatedly dried out in places. Three hundred and thirteen
tracks were recorded on nine levels, including those of five different
dinosaur species. Tracks of ankylosaurs were the first to be discovered
there. The high diversity of tracks clearly shows that the decline of dinosaur
species toward the end of the Cretaceous, at least in South America, was not
gradual, but that the dinosaurs died out abruptly due to external factors.

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 Chapter 12

Birds—The Last Dinosaurs

Where Is the First Ostrich?

The only dinosaurs that survived the mass extinction at the end of the
Cretaceous period are the modern birds. We can divide them into two major
groups: the primitive “old jaw” birds (Paleognathae) and the “new jaw”
birds (Neognathae). The distinguishing features of these two groups are,
among others, the palatal roof on the beak of the birds and their pelvis,
where there is an opening between the ischium and the hip bone in the
paleognaths. This opening in paleognaths is clearly reminiscent of other
non-avian dinosaurs. In neognath birds, the skeleton is already further
derived and differs more from the original dinosaur bauplan.
The paleognath birds include, for example, ostriches, cassowaries, emus,
and the South American nandus, which I saw in the wild during my
excavation trip to Argentina. The extinct moa and the elephant bird belong
to this group also. The surprise representative of the group is the kiwi,
which is closely related to the cassowary. The only paleognaths that can fly
are the tinamous. Like the nandus, they originate from South and Central
America and are found in the tropical lowlands. All other birds belong to
the subclass of neognaths, which can be further divided into the
Galloanserae and the Neoaves. The word Galloanserae sounds complicated
and hard to remember, but in reality it is just a composition of the two Latin
words gallus, which means “chicken,” and anser, the Latin word for
“goose.” Accordingly, this group includes all galliformes and all waterfowl.
The galliformes include pheasants, quail, and guinea fowl, and the
waterfowl include geese, ducks, and swans. The name Neoaves simply
means “the new birds.” They include 95 percent of all birds alive today,
which is still over ten thousand species. By contrast, the paleognaths, i.e.,
all ratites and tinamous, together comprise only about sixty extant species.
But although their diversity is very minimal compared to the new birds,
they occur on three continents and play important roles in their respective
ecosystems. Ostriches and their close relatives are quite poor at
distinguishing colors. This is an important difference from the neognath
birds. I will explain why this is significant when I write later about color
vision in dinosaurs and birds. All of the paleognaths lay their eggs in nests
on the ground, with the rooster performing many of the brood care tasks
and helping the hen hatch the eggs. The eggs of this group are the largest in
the animal kingdom, led by those of the ostrich. However, the eggs of the
extinct elephant bird were much larger. The content of one such egg was
equivalent to the mass of about one hundred and ninety chicken eggs. The
egg of the kiwi is much smaller, but this bird lays the largest eggs relative to
its body size. A kiwi egg can weigh up to five hundred grams, which is
about 30 percent of the mother’s body weight. In this case, the egg fills
almost the entire abdominal region of the hen. The size of these eggs can be
explained by the fact that paleognath chicks are precocial and already very
well developed by the time they hatch. Their bodies are more developed
than those of altricial chicks even when they have just hatched. As a result,
the chicks are also larger and often already have plumage, whereas some
altricial chicks are still naked and blind at hatching. Although it can be
advantageous to give birth to young that are already well developed and no
longer need intensive care from their parents, chicks cannot remain in the
egg indefinitely. The size of the eggs of ostriches, moas, and elephant birds
is already approaching biological limits. Even dinosaurs that weighed ten or
even a hundred times more than ostriches could not lay larger eggs. The egg
of the largest moa, Dinornis, is about as big as that of a Tyrannosaurus rex,
although the Tyrannosaurus weighed about fifty times more than Dinornis.
But why is the T. rex egg not larger? Why can eggs not grow to any size?
As we all know, the shell of bird eggs is made of calcium carbonate
(CaCO3) and is very porous to allow the embryo to breathe air in and for
carbon dioxide to escape when it exhales. With a larger egg, the thickness
of the shell would also have to increase to keep the egg stable and prevent it
from being crushed by its own weight or that of the incubating parents. But
the thicker the shell, the less air gets inside, and the less carbon dioxide
(CO2) can escape; ultimately the unhatched chicks would suffocate. In the
case of the ostrich, this is exactly what happens during the last phase of
growth in the egg: the chick literally loses its breath. And when it then
convulses and begins to twitch inside the egg as a result of its impending
death from suffocation, and due to the violent spasms and kicks of the chick
inside, the thick eggshell finally breaks. Male ostriches in the wild weigh
about two hundred and forty pounds and females slightly less. In the first
year of life they gain about two hundred and twenty pounds in body mass—
so they grow incredibly fast, which is only possible due to their active
metabolism. To do this, they need a lot of food, but also a lot of oxygen.
However, while the surface of their egg grows according to a quadratic
function, the volume inside increases to the third power. So, if the egg grew
any larger, the ratio of surface area to volume would change so unfavorably
that not enough air could get inside and the chick would suffocate, since the
surface of the eggshell is its only access to atmospheric oxygen.
Conversely, as the size of the embryo increases, so does the amount of
carbon dioxide that accumulates inside the egg. If the shell becomes too
thick and carbon dioxide is exhaled regularly at the same time, it can no
longer escape quickly enough. The same is true for dinosaurs. Today we
know that these prehistoric animals, at least in their growth phase, had an
active metabolism comparable to that of birds. They, too, grew very
quickly, and they, too, would have run out of air in even larger eggs.
But paleognaths are fascinating not only because of their unusually large
eggs. Their feet strongly resemble those of large predatory dinosaurs.
Ostriches, nandus, cassowaries, and emus are also much larger and
significantly heavier than any birds capable of flight. The giant Kori
bustard, weighing up to forty-two pounds, is the heaviest bird capable of
flight. But even the nandu, the smallest ratite, already weighs about fifty
pounds. Emus weigh up to eighty-two pounds, cassowaries about ninety-
seven pounds, and the African ostrich weighs up to two hundred and forty
pounds—ten times heavier than a swan. Although ostriches are all
vegetarians, some species can be very aggressive and dangerous. There are
several documented cases where cassowaries have fatally injured humans
with their sharp claws. It is hard to imagine how much worse the T. rex’s
attacks must have been, considering it weighed two hundred times more.
The similarity of the paleognaths to the large predatory dinosaurs
suggests that this group of modern birds arose earlier than the neognaths.
Some scientific studies involving genetic data and paleontological values
indicated that all modern orders of birds, including the paleognaths, must
have originated in the Early Cretaceous and even then began to branch off
from the phylogenetic tree of the giant theropods. Molecular data suggest
that the paleognaths evolved as early as 110 to 120 million years ago in the
Lower Cretaceous, just as the first enantiornithines did. Unfortunately, we
have no fossils from that time. The oldest unequivocal evidence for a
paleognath in the fossil record comes from strata that formed only after the
impact of the asteroid. The oldest member of the group known to date is
called Lithornis celetius and comes from rocks from the Middle Paleocene
of Montana and Wyoming. It is about 61 to 60 million years old. But where
are its older relatives?
If paleognaths were formed 110 to 120 million years ago, but the first
fossils do not appear until the Paleocene, this leaves a gap of about 50
million years in the fossil record, which is hard to explain. When there is
such a large gap between the hypothetical appearance and the first actual
fossils of a group, it is called a ghost lineage. We know that these animals
should be found somewhere, but we have no trace of them. Perhaps older
specimens will indeed be found one day, closing the gap. But maybe the
molecular data has led us down the wrong track. In the meantime, there is
growing evidence that the diversification and evolution of these animals
accelerated after the asteroid impact. It is possible that we are dealing with
a phenomenon similar to the rapid diversification of ichthyosaurs at the
beginning of the Triassic. Perhaps the speed of molecular evolution was
underestimated and perhaps the evolution of birds—just like in ichthyosaurs
—accelerated after the impact of the asteroid as a result of adaptation to
new habitats. The last common ancestor of all present-day paleognaths
could also have arisen just before the end of the Cretaceous, surviving the
extinction event and only then evolving from a small form to a large,
flightless animal. Unusually high mutation rates would have ensured rapid
diversification.
However, the question of the first appearance and origin of the group is
not the only unsolved mystery in this context. Today’s paleognaths all come
from countries in the southern hemisphere that once belonged to the
continent of Gondwana.
 The large continent Gondwana consisted of today’s South
America, Africa, India, Australia, and Antarctica.

Therefore, it was long assumed that this group originated there. However,
molecular data, the fossil record, and phylogenetic studies over the last
twenty years refute this hypothesis. Animals such as Lithornis also suggest
that the evolution to flightlessness and large body sizes occurred later in
their evolution. So, the question arises whether it is possible that these two
traits evolved three times, independently of each other, in South America,
Africa, and New Zealand, at a time when these landmasses were already far
apart. Thus, we still do not know exactly how, when, and where the present
paleognath diversity arose. The group does appear to have lived before the
Cretaceous–Paleogene boundary (K–Pg), but since we do not have
Cretaceous fossils, we cannot trace the convergent transitions to
flightlessness and large body sizes. And while Lithornis is about 60 million
years old, we don’t even find direct ancestors of modern paleognaths until
about 40 million years later.
Moreover, the following thought is keeping me up at night: paleognaths
and neognaths are the only surviving dinosaurs. In the group of archosaurs,
only the modern crocodiles have survived. This means that the closest
relatives of the paleognaths outside the avian lineage are the crocodiles.
This split of archosaurs into crocodiles and dinosaurs, however, already
occurred 250 million years ago. So, if we compare features in the family
tree or in the genome, it is not surprising that all paleognaths always end up
in the same group. They are different from the other birds and are light-
years away from crocodiles in their development. But maybe this effect is
just related to the fact that we don’t have any dinosaur DNA available yet?
Maybe the paleognaths are not a uniform group at all, and their similar
appearance is only a result of convergent evolution? Perhaps the
bioprovince of Australia was already so isolated that cassowaries and kiwis
descended from a different coelurosaur than ostriches, nandus, and
tinamous. Or maybe it all happened much later. We can only hope that at
some point there will be finds that will allow us to reconsider the
relationships of paleognaths to neognaths and their dinosaurian ancestors.
This would also explain the long ghost lineage, because there is no direct
common ancestor. Perhaps ostriches, nandus, and cassowaries split off from
coelurosaurs at three different nodes.

Almost Like Easter:

An Egg Thief Who Lays Colorful Eggs

All birds lay hard-shelled eggs. They are also the only terrestrial
vertebrates, some of which lay colorful eggs. Crocodiles and turtles are a
little different. They also lay eggs, but these are white because their shells
do not have color pigments. From whom did birds inherit the ability to lay
colored eggs? Were their dinosaurian ancestors also able to do this? Did all
dinosaurs lay colored eggs? And why do birds lay colored eggs while
crocodiles do not?
That some birds produce colored eggshells is a result of selective
pressure. Egg coloration is primarily for camouflage. If they have the same
coloration as the vegetation that surrounds them, or if they have speckles to
blend in with gravelly substrate, they are less likely to be discovered, and
are less likely to be eaten by egg thieves. Egg camouflage is thus an
important part of the breeding behavior of open-nesting birds, which protect
their unhatched offspring from visually oriented predators in this way. But
not all eggs of open-nesting birds are colored. In fact, many birds protect
their eggs by continuously incubating, so coloring is not necessary for them.
Colored eggs also help birds distinguish their own eggs from those of other
species, and it keeps them from mistaking foreign eggs as their own, when a
parasite secretly adds them to the nest, as is the case with cuckoos. Turtles,
on the other hand, bury their eggs in the sand, and crocodiles cover them
with branches or twigs, so they don’t need camouflage. If the egg of an
extinct animal is colored, then we can conclude that it was not buried, but
laid in an open nest. The color of its egg thus allows us to draw conclusions
about the social behavior of an animal, even if we can no longer observe it
in the wild. This has led dinosaur researchers to investigate whether
dinosaurs laid colorful eggs even before modern birds emerged from them.
Fortunately, there are many sites with dinosaur eggs around the world. The
first specimens were discovered in France in 1846. Other finds have come
from the United States, Spain, and China, where entire nests and clutches
have been found containing dozens of eggs.
 Jasmina Wiemann, a colleague at the Field Museum, wanted to know if
only the direct ancestors of birds produced colorful eggshells or perhaps
long-necked dinosaurs, duckbill dinosaurs, or horned dinosaurs as well. She
studied dinosaur eggshells from around the world and used
microspectroscopy to identify different pigments responsible for bluish,
greenish, and reddish-brown coloration. She found that color pigments are
present only in the eggshells of theropods, but not in other dinosaurs. She
also found that among carnivorous dinosaurs, only Maniraptora had colored
eggs. Carnivores such as Tyrannosaurus and Spinosaurus did not lay
colored eggs.

Maniraptora means “Seizing Hands.” These dinosaurs

evolved a crescent-shaped wrist bone that allowed them
greater hand mobility, which was a prerequisite for bird
flight. Feathers also evolved within the Maniraptora group. In
the Mesozoic, these animals tended to be small-to medium-
sized and had the largest brains among dinosaurs relative to
their body weight. The group also includes modern birds.

Jasmina Wiemann was able to trace the origin of colored eggshells back
to an oviraptor from the Late Cretaceous of China. Oviraptors are those
toothless maniraptors we know as “egg thieves.” Yet they were probably
very caring parents, hatching their chicks in their nest and nurturing their
young. Their nests contain eggshells of the type Macroolithus yaotunensis,
assigned to the species Heyuannia huangi. Some of these eggs contained
exceptionally well-preserved embryo remains that can be assigned to
Heyuannia. They come from three river deposits from eastern and
southernmost China. The pigments in these eggshells indicate a blue-green
coloration. The colored eggs of Heyuannia are thus the oldest in the fossil
record. Their pigmentation supports the interpretation of their depositional
conditions—because the eggs were colored, they were laid in an open nest
and were not buried. The nests of these oviraptors are mostly circular and
the eggs in them are arranged in several concentric layers, stacked on top of
each other. All the while, the elongated eggs are stuck almost vertically in
the sediment, with the pointier side down. In the circle, they are also
arranged in pairs, with two eggs always close together and separated from
the next pair by sediment. This arrangement, as well as the shell pattern and
shell porosity, are clear indications of an open nesting behavior of
Heyuannia, because buried eggs, as those of turtles, for example, are not
neatly, symmetrically arranged.

It is interesting to note that no bird today arranges its eggs in this way.
Only in the nests of ratites, which live in breeding colonies, can a similar
breeding behavior be observed. Some of these birds lay green ones, like
emus, or bluish ones, such as cassowaries. The nest arrangement and blue-
green eggs indicate that oviraptors were already engaged in intensive
nesting. The colors blue or green are usually not used for camouflage, but
as signal colors. They are especially common in birds, whose roosters play
a major role in parental care. So, we can learn and infer a lot about the
social behavior of these animals from the color of their eggshells.
Because we have not found colored eggshells in other dinosaur groups,
the blue-green eggs of Heyuannia also show that colored eggshells did not
evolve multiple times, but only once in the Maniraptora group. So, if
someone asks you in the future who brings the colorful Easter eggs, the
answer should definitely be the Maniraptora!

The Goddess Asteria:

The Mother of All Chickens and Geese

The origin of modern birds remains difficult to find, as shown by the ghost
lineage of paleognaths. We cannot trace the earliest evolutionary stages of
modern birds because of the incompleteness of the fossil record from the
Mesozoic. Phylogenetic analyses suggest that modern birds split off in the
Cretaceous, but representatives from the ancestral lineage of birds are
virtually unknown from the Mesozoic. The first paleognaths do not appear
in the fossil record until about 60 million years ago, although they are
actually thought to have existed much earlier. The situation is similar for the
neognath birds. Key questions about their geographic distribution and
ecology, and the actual divergence of modern birds, therefore remain
unanswered. However, a Mesozoic fossil was recently discovered in
Belgium that is undoubtedly a modern bird from the neognath group. The
fossil fills an important phylogenetic gap in the early evolutionary history
of the so-called crown birds. Some characteristic features of its skull
indicate that the animal was closely related to the last common ancestor of
chickens and geese and may be the last common ancestor of Galliformes
and waterfowl—and hence, the first known representative of the
Galloanserae. The find is from the latest Upper Cretaceous period and is
about 66.8 to 66.7 million years old. Hence, it lived contemporaneously
with Tyrannosaurus and Triceratops, about seven hundred thousand or
eight hundred thousand years before the asteroid hit. My doctoral advisor,
Daniel Field, studied it with scientists from Cambridge, Maastricht, and
Greenwich. The fossil consists of an almost complete, three-dimensionally
preserved skull and individual elements of the postcranial skeleton. This
makes it the first modern bird from the Mesozoic with a well-preserved
skull. It represents one of the few unequivocal pieces of evidence of crown
birds from the Mesozoic and exhibits a unique combination of galliform
and waterfowl features. Interestingly, it was also found in strata in which
extinct tooth-bearing seabirds appear that do not yet belong to the crown
group: relatives of Ichthyornis. The co-occurrence of this animal with
Ichthyornis relatives may be evidence of their coexistence. It also
challenges previous hypotheses that crown group birds originated in the
southern hemisphere. The team named the small bird Asteriornis
maastrichtensis. The genus name means “Asteria’s bird,” while the species
name refers to where it was found and its geological age. Both names take
into account other additional aspects of the fossil. Namely, Asteria was a
Titan from Greek mythology who transformed herself into a quail to escape
Zeus’s advances. Quails are galliform birds and Asteriornis is the mother of
all galliformes. Also, Asteria once plunged from the sky into the sea—just
like the asteroid that fell into the sea off the coast of Chicxulub at the end of
the Cretaceous period.
The importance of this find can hardly be overstated: first, because it
provides us with new information about the habitat of the first modern
birds; and second, because finds of the very first modern birds are
extremely rare. In fact, there is only one other bird from the late Upper
Cretaceous that undoubtedly belongs to the modern birds and of which
more than a single isolated bone has been found. This bird is called Vegavis
and is about 66.5 million years old, or about two hundred thousand to three
hundred thousand years younger than Asteriornis. It was found on Vega
Island, which is off the Antarctic Peninsula. This is where the name of the
animal comes from. Some colleagues suggest that Vegavis is closely related
to ducks and geese, while others believe it is still phylogenetically just
outside the Galloanserae. To be certain of the answer to this question, we
would need further skull material of the animal, through which we could
recognize characteristic features. However, there is more evidence that it
belonged to the neognaths than it being an ancestor to the group. In any
case, Asteriornis and Vegavis have relatively small body sizes, and both
finds come from sediments that indicate they were coastal dwellers. They
lived in a similar ecosystem, which may provide an explanation for why
modern birds were able to survive the asteroid impact. I return to this
question in the last chapter of the book.

Beyond the Rainbow

Although we do not know exactly when the first feathers emerged, it is

clear that some dinosaurs possessed them—long before birds could take to
the skies with their help. So, feathers are not a unique feature of birds, and
they have served many other purposes before bird flight. In modern birds,
contour feathers form the majority of the visible plumage. The contour
feathers covering the body are called the pennaceous feathers.
Those pennaceous feathers that are used for flying are fittingly called
flight feathers. They are long, stiff, asymmetrically shaped, but
symmetrically paired on the wings or the tail of the bird. The flight feathers
on the wings are called remiges (singular remex), and those on the tail are
called rectrices (singular rectrix).
 In addition to the contour feathers, birds have down feathers that insulate
the body. Their pennaceous feathers are necessary for flight, but also for
communication, camouflage, and breeding. Which function the pennaceous
feathers originally served in dinosaurs has long been unclear. We don’t
know exactly when dinosaurs first developed contour feathers; the oldest
evidence comes from the Jurassic period. The remains of some
Archaeopteryx specimens show that these animals also had contour feathers
about 150 million years ago. Closely related to Archaeopteryx and even
older is a fossil from Hebei Province in China, east of Beijing. It is about
161 million years old and has asymmetrical contour feathers. Whether or
not Archaeopteryx was able to fly actively is still a matter of debate.
However, it is undisputed that it was able to glide and flutter. This is shown
by its almost modern-looking asymmetrical flight feathers and the clavicles
which are fused into one wishbone. In an asymmetrical feather, the vanes
on the left and right of the quill are of different widths. Such asymmetry is
considered a mandatory requirement for powered flight. It gives the birds
aerodynamic characteristics that are essential for flight. The strong
wishbone also indicates that the animal’s thoracic muscles were particularly
robust and that the animal could perform powerful strokes with its wings.
Whether this enabled Archaeopteryx to lift off the ground under its own
power cannot be said with certainty. The backward-facing first toe of its
foot could be evidence that it was an arboreal dweller that used this claw to
climb or cling to branches. The trees may have served as a launching pad
for when it was unable to get off the ground. However, the area of the
Solnhofen Limestone in Bavaria, where these proto-birds (Urvögel) were
found, are deposits of a lagoonal landscape that formed on the edge of the
Jurassic Sea in a hot, dry climate, where there were few or perhaps no trees.
Archaeopteryx would have had to climb high cliffs and swoop down from
them to demonstrate its flying skills. In the eleventh specimen of
Archaeopteryx discovered so far, some of my colleagues from Bavaria have
discovered spectacular plumage that covered its entire body, even its legs,
with long feathers, arranged in rows. In modern birds of prey, such long leg
feathers do not actively contribute to flight but are helpful when landing.
According to Christian Foth of the University of Fribourg, who studied this
specimen with his team, the feathers may have not only served for flight,
but mainly for display because of their smooth surface and light-refracting
properties. Thus, contour feathers were superior to filamentous
protofeathers in courtship. Protofeathers probably evolved in early
dinosaurs. Initially, their main function was exclusively thermal insulation.
Thus, they contributed to an increase in growth rate, which was
accompanied by a faster metabolism at the beginning of dinosaurian
evolution. This requires effective insulation of the body; otherwise,
metabolic energy is lost as heat instead of being available for growth. This
was especially crucial for small-to medium-sized early dinosaurs and their
young, which had an unfavorable mass-to-surface ratio. While the long-
necked dinosaurs and the tyrannosaurids grew larger and larger over the
course of their evolution, the representatives of the avian lineage began a
reduction in body size. This miniaturization with simultaneous rapid growth
and high metabolism was only possible due to good body insulation.
Now it was the case that the dinosaurs had already developed colorful
scales to attract mates or deter rivals. These scales must have been very
colorful and arranged in species-specific patterns. The special arrangement
of scales of different sizes suggests that these scales also had different
colors. Therefore, dinosaurs may have had stripes like tigers or zebras, or
spots like leopards, and at the same time exhibited such a beautiful play of
colors as sand lizards. However, this blaze of color was limited by the first
protofeathers—namely, the hairlike filaments entailed the loss of structural,
color signaling capabilities. This is still the case today in birds, whose body
coverings consist of flexible, filamentous feathers. Iridescent, variegated
colors are only possible in tropical birds because they have contour
feathers. Hair or protofeathers cannot produce such color effects.
Displaying iridescent greens and blues or ultraviolet color reflections
requires a precise arrangement of nanometer-scale light-scattering elements,
which in modern bird feathers consist of keratin, melanosomes, and even
air. It is through the tiny but highly ordered rami and barbules in vanes,
interlocked by tiny hooks, that a bird produces the iridescence and rich play
of colors of its contour feathers. These colors are essential for
communication and sexual selection. Thus, to maintain the rich blaze of
color of their scales, tiny dinosaurs could not simply rely on protofeathers,
but had to evolve more complex feathers that included keratin,
melanosomes, and air in their structure. At the same time, the colorfulness
of feathers is only as good as the visual ability of their counterparts allows
—which raises the question of the visual color spectrum of dinosaurs. We
know that virtually all of today’s reptiles can perceive colors in a much
more sophisticated way than humans and other mammals. Many reptile
species can even see ultraviolet colors. We are not able to do that. If we
infer phylogenetically, this suggests that dinosaurs were also equipped with
the highly differentiated color vision of birds, known as tetrachromacy.

Cone cells in the retina of the eye are responsible for the
perception of colors. Cone cells are neurons that are needed
as specialized sensory cells for seeing in daylight. Mammals
have three different cone cells and are therefore called
trichromats. Reptiles have four different cones, which is why
they are called tetrachromats. Trichromats can recognize
about a million different colors, and tetrachromats up to a
hundred times more. It is believed that they can distinguish
about 100 million colors. Three of their cone cells perceive
the colors blue, green, and red, and via a fourth, short-wave
receptive neuron they can recognize the ultraviolet color
spectrum up to shades of turquoise.

Thus, pennaceous feathers may have solved another very complex

evolutionary problem in addition to enabling powered flight. Maniraptors,
for example, independently evolved feathers in different branches that were
clearly not for flight but for good insulation, to facilitate increased
metabolic rate, and miniaturization without losing the structural color
signaling of their scales. This would not have happened with a complete
covering of protofeathers or hairs. This is because fur, unlike feathers, has a
limited color spectrum, because without a coherent surface it does not
enable directional light scattering and compromises structural color
representation. The color pigments in hair therefore produce only reddish-
brown to black tones, with lighter colors achieved by the absence of
pigments.
In mammals, too, there was a trend toward reducing body size and
increased thermal insulation through thick fur. However, because mammals
were nocturnal in the Mesozoic era, sophisticated color perception did not
play a role. At night, all cats are gray. Tetrachromacy is probably an original
feature of terrestrial vertebrates, and in the ancestors of modern mammals
differentiated color vision was lost because it had no meaning at night and
played no role in their survival.
In dinosaurs, however, flat or planar structures gradually evolved from
hairlike feathers. Planar feathers initially obscured the filamentous
protofeathers and replaced them over the course of evolution, eventually
leading to the development of smooth contour feathers. And while we do
not know exactly when the first protofeathers appeared, it seems at least
clear that the first planar contour feathers occurred in the Maniraptora
group. In them, the rami and barbules of the contour feathers were in one
plane. The hooks and barbs, with which the barbules were held together,
could, just like in a zipper, be opened and closed as often as desired. This
made the feathers extremely durable despite their lightweight construction.
In addition, the fact that the rami and barbules were positioned closely next
to each other on a single plane created a particularly smooth, flat, and
uniform surface. This was particularly important because the precise
arrangement of the light-scattering pigments allowed optimum light
penetration, which was a prerequisite for the iridescence of the feathers.
The birds achieve the best color play only with these flattened filaments.
The smooth, foil-like surfaces provide optimal physical conditions for the
iridescent rainbow colors. It is surely no coincidence that the maniraptors,
of all dinosaurs, who had very mobile wrists, also had large feathers on
their arms and hands that allowed them to maximize surface area and create
structural color signaling on a macroscopic level. Large, smooth feather
surfaces acted as a canvas for detailed ornamentation and patterns.
Ostriches, which have frizzy plumage, as opposed to a well-structured
planar plumage, have only brown, black, or white integuments (just like
mammals), and cannot produce the complex variety of colors seen in
contour feathers. If we remember that paleognaths are the only modern
birds that distinguish colors poorly, this also makes sense. A special case
among paleognaths is the helmeted cassowary, which can probably see
more colors than its relatives. The necks of cassowaries shine a bright blue
that comes not from the feathers, but from the strong skin color on the neck.
The fact that paleognaths are the only birds today that have filamentous
feathers should definitely be given greater consideration in future
phylogenetic analyses. Might it be possible that they split off from
coelurosaur ancestors even before the Maniraptora evolved? If they
underwent convergent evolution, would we recognize that they merely
resembled the Maniraptora and other present-day birds? Would we
recognize that they do not have the same maniraptoran ancestor as the
neognath birds—and that even the paleognaths are descended from different
coelurosaurs, when their closest relative outside of birds is the crocodile?
In this context, the compsognathids, a group of very small carnivorous
coelurosaurs, have caused a stir in research circles. In a compsognathid
from China, a lush, fur-like plumage was discovered. Fossil melanin was
found in the compsognathid Sinosauropteryx, whose name means “Chinese
reptile wing.” The melanin allowed its plumage color to be determined.
Lacking planar feathers, this animal did not have a wide range of iridescent
colors. Instead, during its lifetime, its fibrous feathers were brown to
reddish brown in some places and white in other places. What made this
find special, however, was not the colors alone, but a countershading, as we
have already encountered in the armored Borealopelta. Sinosauropteryx
comes from the Early Cretaceous strata of the Jehol ecosystem in
northeastern China, which was previously thought to have consisted of
dense, closed forest. Characteristic color patterns in the plumage of the
dinosaurs that lived there may confirm this, because light conditions in the
forest were different from those in open terrain. However, the coloration of
Sinosauropteryx does not indicate a forest dweller, but rather an
unprotected, wide-open habitat. To be sure that the colors of
Sinosauropteryx were indeed fossil melanin, two animals of this species
were examined, and intriguingly, nearly identical color patterns were found
in both. The bridge of the nose, the lower jaw, the throat, and the chest were
white, but the area around the eyes was dark. Thus, Sinosauropteryx looked
almost like a raccoon. However, such “bandit’s masks” are not only typical
for raccoons, Zorro, or the Beagle Boys (from Donald Duck cartoons); they
are a common pattern in many birds, where they serve as camouflage. The
tail of Sinosauropteryx, on the other hand, was very reminiscent of the curly
tail of a lemur, with alternating white and reddish-brown bands. In any case,
color reconstruction helps to gain a better understanding of habitat, and we
can tell more about ecology from contour shading than from skeletal
features alone. The fact that small predatory dinosaurs also lived in open
habitats in the Jehol bioprovince contributes to a clearer picture of the
environment and fauna, and therein may lie the key to the great biodiversity
in the Jehol Basin. The extraordinary species richness of this area can be
more easily explained if we consider that this region has evolved over a
period of more than 10 million years, and that the vegetation, as well as the
landscape, has changed drastically several times during this period. It is
therefore not a contradiction when we find both forest and steppe dwellers
in the Jehol bioprovince, where the most spectacular feathered dinosaurs of
China come from. The biodiversity is indeed remarkable. Here lived not
only enantiornithines, but also even more basal, small dinosaurs, such as
microraptors, which wore feathers but were not yet birds even in the
broader sense. They were descended from the sickle-clawed raptors.
Another small feathered dinosaur is called Confuciusornis. They are also
not yet a modern bird, but are at least already on the evolutionary branch
that led to birds. Some of them have extravagant decorative feathers on
their tails and at least outwardly remind us a little of birds of paradise.
There are hundreds of fossils of these animals, and Chinese researchers
were able to determine a sexual dimorphism in them, where one sex type
had particularly long and extravagant tail feathers and the other did not.

What Microraptor, Confuciusornis, the enantiornithines, and the “almost-

birds” of Jehol have in common are their contour feathers. Unlike
Sinosauropteryx, they all already had smooth, planar pennaceous feathers
and thus probably had iridescent plumage. With their tetrachromatic vision,
they lived in a motley world and could, with the fourth cone cell type in
their retina, perceive a hundred times more colors than we humans. That the
dinosaurs loved iridescent colors is not only proven by the shape of their
feathers. In the case of the Archaeopteryx relative from Hebei Province,
which I mentioned earlier, there was also evidence of melanin. Unlike
Sinosauropteryx, however, it had iridescent colors because its melanin was
discovered in its contour feathers, which is why the animal became known
as the “rainbow dinosaur.” The name of this newly discovered dinosaur
species is Caihong juji, which is Mandarin and means “the rainbow with the
big crest.” This term refers to the animal’s large, broad, and iridescent
feathers and the bony crest it wore above its eyes.
The platelet-shaped melanosomes of Sinosauropteryx were discovered
for the first time under a scanning electron microscope. They were still
intact and could be compared with those of modern birds. The feathers of
Caihong bear a strong resemblance to those of hummingbirds. Their
melanosomes are actually black, but light is refracted by the special platelet
shape to create a shimmering effect. Modern birds use their colorful
plumage to attract mates. Accordingly, Caihong’s rainbow feathers could be
a prehistoric version of a peacock’s iridescent tail. Caihong is the oldest
known example of platelet-shaped melanosomes typically found in bright,
iridescent feathers. It is also the oldest animal known to date with
asymmetrical feathers.

OceanofPDF.com
 Chapter 13

The End of the Dinosaurs

The Day it Rained Glass

On an otherwise uneventful day in spring 66 million years ago, a huge
boulder from space, the size of New York, crashed into Earth. As
fascinating as the dinosaurs, the flying reptiles, and the marine reptiles
were, they were nevertheless wiped out by this asteroid impact at the end of
the Cretaceous period. They did not manage to escape the cosmic
catastrophe. While the birds’ survival also ensured the continuation of
asymmetrical feathers, other innovations of nature were lost forever. Today,
there is no animal that can fly underwater like a plesiosaur with four
paddle-shaped fins. There are no flying animals that have wingspans of
more than thirty or so feet, or long-necked giants that weigh more than fifty
tons. Crocodiles cannot gallop today, and there are no carnivores that weigh
fifty times more than modern lions. Some ecological strategies have also
been lost, such as the splitting of an animal species into different ecological
niches, as we have seen in long-necked dinosaurs and tyrannosaurs. This
strategy, which was incredibly successful over millions of years, ended up
sealing their fate.
Traveling at a speed of fourty-four thousand miles per hour or more, and
with a diameter of more than six miles, the asteroid hit so hard that it even
melted the Earth’s crust, locally. The extraterrestrial object crashed into
Earth off the coast of the Yucatán Peninsula in Mexico. The small village
Chicxulub is the closest human settlement to the impact site, which is why
the asteroid is now called “the Chicxulub impactor.” The impact was
followed by tremendous shock waves that formed tsunamis in the Gulf of
Mexico that were more than a mile high. In Big Bend National Park in
Texas, about a thousand miles from this site, we find sediments that were
formed immediately after the impact and are called tsunamites.

Tsunamites are sedimentary rocks composed of the sediments

formed during a tsunami.

The impact of the asteroid not only caused earthquakes and tsunamis, but
also ejected huge amounts of debris, molten rock, and dust from the crater.
The force of the impact was so violent that some pieces of rock may have
even been hurled into space. At the Tanis Fossil Site in North Dakota, about
two thousand miles from Chicxulub, researchers have found tektites
associated with the asteroid.

Tektites are glass objects formed when surrounding rock

melts during an asteroid or meteorite impact and is hurled
away from the impact site. The mostly teardrop-shaped
objects can grow to several inches in size and are often
composed of high-pressure modifications of quartz.

In the Tanis sediments, which are part of the Hell Creek Formation, fish
skeletons were found that had been hurled ashore and perforated by small
tektites. In the fossilized remains of some of these fish, the small projectiles
were still stuck in their gills. The fish were an important indication of the
seasonal confinement of the impact event. Indeed, the earthquake deposits
show an annual cyclicity in the last years of the Cretaceous and prove that
the impact occurred during the northern hemisphere’s springtime.
I remember the early phase of the research project surrounding the Tanis
fossil site well—I was not involved myself, but was lucky enough to meet
Melanie During, the lead author of the paper, in Grenoble in 2018. I was
scanning Mesozoic and recent fish with our Oxford team while Melanie
was there segmenting fish fossils and the small, spherical tektites for her
project. I had seen her at a few meetings, but I had never talked to her
before. In Grenoble, she took the time to explain her work to me, which she
published in Nature in 2022. I found the topic exciting at the time, not
realizing that her research would later receive so much attention. As it
turned out, she could detect seasonal changes in the bones of the fish, just
as we have seen in the bones of Thrinaxodon. Just like Thrinaxodon, there
were signs of aestivation and hibernation in the fish. In addition, their
reproduction apparently followed annual cycles, and the food supply also
varied depending on the season, which of course makes sense, since
otherwise aestivation and hibernation would not have been necessary.
It is amazing that the Tanis site provides an accurate chronology of the
immediate events of the Chicxulub impact. The strata there were formed
within about an hour of the impact and represent—far inland—a mixture of
marine and terrestrial deposits. Although the nature of their assemblage is
reminiscent of tsunamites, these layers were likely formed by the shock
waves of a massive earthquake. Researchers estimate the earthquake that
could lead to such strata had a magnitude between 10 and 10.6 on the
moment magnitude scale (MW). A 10 MW earthquake already releases the
energy of 1.2 million Hiroshima bombs. The most violent earthquake ever
measured in our time occurred in Chile in 1960 with 9.5 MW. At first, this
may sound similarly devastating as the primeval asteroid impact; however,
an earthquake with a magnitude of 10.5 MW is about thirty-six times more
powerful than one with 9.5 MW. The moment magnitude scale ends at the
value of 10.6 MW, because physicists assume that the Earth’s crust would
break apart completely. So, if the asteroid had been just a little bit bigger
and crashed into Earth a little bit faster, it might have destroyed the whole
planet.
Evidence of the dust cloud caused by this impact, which completely
enveloped the Earth for a short time, are wafer-thin layers of sediment
known as the iridium anomaly. In 1980, this anomaly, detectable around the
world, was scientifically described by Luis and Walter Alvarez, Frank
Asaro, and Helen Michel. The team was able to detect an elevated
concentration of iridium in a boundary layer between Cretaceous and
Paleocene rock deposits in Italy and Denmark. Such layers have since been
detected in New Mexico, New Jersey, and other regions. In 2022, an article
was published that reported on rock layers at the Cretaceous–Tertiary
boundary from Baja California, Mexico. They consist of terrestrial and
shallow marine sediments that were rapidly rebedded onto sediments on the
continental slope as a result of the impact. Radiometric dating indicated that
the strata are about 66.12 million years old, with a margin of error of
650,000 years. This corresponds to the age of the Cretaceous–Paleogene
boundary. They contain corals, marine gastropods, and bivalves, as well as
tuffs, quartz, and charred logs that originated on land. The quartz grains are
shocked quartz, formed only when earthquake waves pass through them.
This unusual mixture of disparate materials is interpreted as heterogeneous
landslide deposits generated by an earthquake as a direct result of the
Chicxulub impact and a mega-tsunami. The charred logs in the sediments
probably formed in a very short period of time. Immediately after the
impact, a huge fire wave with a temperature of more than 1,832°F must
have swept over the land at this location, which was extinguished by the
tsunami only a few minutes later. As the water flowed back, the floods then
swept away trees, tuffs, and coastal sediments, washing them into deeper
water. Directly above, we now find mudstones that also bear a distinct
iridium signature. While the Tanis sediments were formed within an hour of
the impact, these sediments were deposited within the first ten minutes after
the impact.
Since the element iridium hardly occurs in the Earth’s crust, but is found
in high concentration in meteorites, an iridium-rich layer can be an
unmistakable sign of an extraterrestrial impact. Some astronomers believe
that the origin of this asteroid was in the farthest reaches or just outside our
solar system in the Oort Cloud. Recent evidence suggests that the asteroid
may have broken apart shortly before the impact, and a second part of the
celestial body crashed to Earth off the west coast of Africa.

Like a Phoenix From the Ashes:

How Birds Survived the Catastrophe
When the asteroid hit the Earth 66 million years ago, all dinosaurs except
modern birds became extinct, as well as the pterosaurs, all marine reptiles,
and the ammonites. The asteroid impact also had a massive impact on plant
life. Several factors caused global deforestation. The shock waves that went
around the world immediately after the impact certainly caused significant
forest loss, and subsequent forest fires devastated large areas of vegetation.
In addition to the immediate effects of the impact, there were long-lasting
factors that damaged the forest. Even in the months that followed, ash,
smoke, and soot in the atmosphere caused reduced solar radiation, which
massively impaired plant photosynthesis while causing global cooling. The
entire world was in a permanent dim state. In the twilight, forests continued
to die. Even as the dust slowly settled, low sunlight and cooler temperatures
delayed recovery in massively damaged forests. Why forest dieback and
ecological collapse were less severe for birds than in their closest relatives
is still the subject of extensive debate. And as simple as the question of why
the birds survived may sound, the answer seems very complex. There are
two ways to ask this question. We can ask once why all other dinosaurs
could not recover from the asteroid impact or in what way the birds differed
from the other dinosaurs that allowed them to survive.
To answer the question of survival, we must first know when exactly
modern birds evolved. The long ghost lineage among ostrich relatives
makes this surprisingly difficult. The first representatives of the group may
have evolved more than 100 million years ago, but we find fossil evidence
of the group only shortly before the Cretaceous–Paleogene boundary. The
Galliformes and waterfowl originated only a few hundred thousand years
before the Chicxulub impact, and all other birds probably thereafter. But
there is hardly any fossil evidence of the early representatives of modern
birds. This is partly because birds are usually very small animals with
delicate bones and are often not preserved in the fossil record. Moreover,
they only play a very minor role in ecosystems at the end of the Cretaceous,
yet.
The first reliable evidence of a true Neoaves is dated to 62.5 million
years ago—that’s about 3.5 million years after the asteroid impact. The
fossil came from the state of New Mexico and was named Tsidiiyazhi abini,
a Navajo term meaning “the little morning bird.” This animal is believed to
be related to today’s mousebirds, which are found exclusively in Africa,
south of the Sahara. This complicates our question, since mousebirds are
closely related to passerines, and genetic studies suggest that mousebirds
and passerines must actually have evolved later than many other Neoaves.
Most phylogenies suggest that flamingos and pigeons were the first “new
birds.” Then came hawks, hummingbirds, cranes, gulls, penguins, and many
others. The core land birds, which include the mousebirds and passerines,
were probably the last group of Neoaves. If we look more closely at the
earliest Cretaceous fossil record, we find that Galliformes and waterfowl
lived in very different habitats than the first Neoaves Tsidiiyazhi.
Asteriornis and Vegavis were more coastal dwellers. And the first known
waterfowl, just like Vegavis, comes from Antarctica. The animal is called
Conflicto antarcticus and was deposited in strata of the López de Bertodano
Formation, dating from the Late Cretaceous to the earliest Paleocene. The
layers in which it was found indicate that the bird lived shortly after the
asteroid impact. But why were these early modern shorebirds or waterbirds
able to survive while the ichthyornithids and enantiornithines were not?
One of the reasons surely lies in the fact that a number of their
characteristics proved to be selectively advantageous after the mass
extinction. These included the lower energy requirements of the birds.
Because of their small body size, they also had lower energy requirements
than most other dinosaurs, while showing a certain adaptability and
flexibility in their diet. With the collapse of many ecosystems, considerable
food resources disappeared, and the earliest birds had to rapidly change
their dietary habits. Perhaps an advanced digestive system helped them do
this, allowing them to switch their diet to insects and seeds. For the small
birds at that time, just a few insects or a few seeds a day were probably
enough to survive. Larger animals, on the other hand, did not find enough
food. The ichthyornithids, which are most closely related to today’s birds,
were much larger and probably predominantly fish eaters. But the fish
faunas were severely damaged by the impact, so it became increasingly
difficult for these animals to find enough food. At the same time, their
energy requirements were higher because they were larger and heavier, and
their specialized fish diet did not allow them to switch quickly enough to
seeds and insects. In the case of enantiornithines, their continued existence
was probably affected by the destruction of forests. They could not
adequately protect and feed their chicks, which were predominantly
precocial.
The other flightless dinosaurs could not survive because of their high
energy demands. However, were the conditions really equally bad
everywhere in the world? Were there not perhaps still isolated small island
populations which had been spared by the catastrophe? Did these islands
maybe also offer somewhat better conditions after the impact? We have
already indirectly addressed the answer to these questions several times.
Dinosaurs were very well adapted to their environment, and in a stable
ecosystem they could excellently assert themselves against competitors and
displace them. But it was this successful strategy that caused their demise in
an ecological crisis. In mammals and modern birds, offspring and adults
occupy the same ecological niche, so survival was possible for the young
only if their parents found enough food. In dinosaurs, and especially in
large predatory dinosaurs and long-necked dinosaurs, the different
generations behaved, trophically, like different species. As a reminder, the
different diets of the young and old led to a reduction in niche diversity
because hatchlings, juvenile large predators, and adults functioned as
distinct ecological morphospecies. The same was true of long-necks, whose
young fed on horsetails, while adult sauropods ate leaves from tall trees. If
the young did not find enough food, they did not grow and starved to death
before sexual maturity, and if the adults did not find food, they died before
they could lay eggs.
There were selective advantages for smaller mammals because, like
birds, they often fed on insects. Larger mammalian species could not
survive either. Because non-avian dinosaurs occupied all ecological niches
during their lifetime, none of them could survive.
 But even for small mammals and birds, resources were scarce. When
their hunger eventually became stronger than fear, it drove them to
selectively scavenge dinosaur clutches, which only accelerated their
extinction.

OceanofPDF.com
 Epilogue

This book is intended to give an overview of the incredible discoveries in

paleontology and to show the fascinating findings that my colleagues and I
have made in recent years. Research is active and we are able to use ever
new methods to unravel mysteries that previously seemed unsolvable.
However, the book contains only a tiny sample of what is currently being
researched, and if it finds appeal, I will be happy to tell more in another
book. For the stories I have told, information has flowed together from
different corners of the globe. Whereas fifty years ago research was
dominated by North American and European paleontologists, today the
most exciting discoveries often come from South America, Asia, Africa,
Australia, and Antarctica. We understand dinosaurs better than ever before,
and yet there have never been so many unanswered questions. With every
door we push open, we face new puzzles to unravel. In the process,
understanding the dinosaurs also helps us better understand our own destiny
and perhaps steer it in the right direction for the benefit of our children.
Three of the largest mass extinction events on Earth are related to the
emergence, evolutionary success, and demise of the dinosaurs. Due to a
constantly increasing concentration of greenhouse gases, galloping effects
occurred, first creeping and then accelerating. Ecological collapse was then
always followed by massive species extinction, triggering a trophic cascade.
Birds survived the last catastrophe because they evolved in ecological
niches that were least affected by changing environmental conditions.
Today, humans are destroying bird habitats and food sources and
threatening biodiversity on such a scale that we may be responsible for the
next great mass extinction. If we are not careful, we will complete what the
asteroid failed to do and ensure the extinction of the last dinosaurs on Earth.
But that need not be the case. We know and understand what is happening
around us. Now it is up to us to take the right actions and listen to those
who understand science and offer us solutions to these challenges.
OceanofPDF.com
 Glossary

A
aestivation: a form of summer hibernation in which the metabolism is
lowered and all activity is suspended or shut down altogether as an
adaptation to adverse environmental conditions. It can occur during periods
of heat or drought, and seasonally in arid climates. Aestivating behavior still
exists today in some moth and snail species.

Albian: the last chronostratigraphic stage of the Lower Cretaceous (112.9 to

100.5 million years before present).

Alcmonavis: a prehistoric bird from sediments of the Upper Jurassic of the

Altmühl valley in Bavaria (about 150 million years old).

Allosaurus: a genus of theropod dinosaurs that lived during the Upper

Jurassic of North America and Europe and was one of the largest predators
of its time.

altricial birds: birds that live in their parents’ nest for an extended period of
time after hatching and require brood care.

ammonoidea: a group of extinct cephalopods that lived 407.6 to 66 million

years ago.

Amniotes: a large clade that includes all terrestrial vertebrates, with the
exception of amphibians, that is characterized by the ability to reproduce in
locations outside of the water.

anagenesis: the evolution of a new species that replaces an old species.

Anisian: the first chronostratigraphic stage of the Middle Triassic (247.2 to

242 million years before present).
anoxic (water): fresh or salt water that is entirely depleted of oxygen.

Archosauria: a clade of tetrapods that includes crocodilians, birds,

dinosaurs, and pterosaurs and probably originated in the late Upper
Permian.

Asteriornis: a genus of birds from the Late Cretaceous of Belgium (about

66.8 to 66.7 million years before present), last common ancestor of all
chickens and geese and thus the oldest known neognath bird.

Aves: the scientific name for the class of birds.

Avialae: the group of theropods more closely related to birds than to

deinonychosaurs.

Baryonyx: a dinosaur from the family Spinosauridae, from the Lower

Cretaceous of England (about 130 to 125 million years before present).

biostratigraphy: the correlation and relative chronological classification of

rock strata based on fossils.

bioturbation: the burrowing of soils and sediments by burrowing organisms.

bone beds: an accumulation of fossils in certain rock layers.

Bone Wars: a term used in the American press and popular science literature
to describe the dispute between two American paleontologists, Edward
Drinker Cope and Othniel Charles Marsh, that lasted from 1877 to 1892.

Borealopelta: a genus of armored dinosaurs from the Lower Cretaceous of

Canada (112 to 110 million years before present), whose skeleton is the
most complete ever found of this group.

Brachiosaurus: a genus of long-necked dinosaur from the Upper Jurassic of

North America and Africa (154 to 145 million years before present) found
in the Morrison and Tendaguru formations.
Brontosaurus: a genus of long-necked dinosaur from the Upper Jurassic of
North America found in the Morrison Formation.

Broomistega: a genus of amphibians from the Early Triassic of South

Africa.

Caihong: a genus within the group Avialae.

Camarasaurus: a genus of long-necked dinosaur from the Upper Jurassic of

North America found in the Morrison Formation.

Ceratopsia: the group that comprises all horned dinosaurs.

Chicxulub: a municipality in Mexico, near the impact site of the asteroid

that hit Earth 66 million years ago.

Coelurosauria: a clade within Theropoda.

cladogenesis: a phenomenon whereby an original species splits into two

new species.

computed tomography: an imaging procedure in radiology that can be used

to x-ray bones and fossils.

concretion: an often round or spherical mineral composite. It consists of a

hard, fine-grained sediment that has been baked by pore water and grown
outward from a crystallization center over time.

confidence interval: an interval in statistics used to indicate the accuracy of

the location of a parameter. The confidence interval indicates the range that
includes with a certain probability the parameter of a distribution of a
random variable. A confidence level of 95 percent is often specified by
researchers to measure the informative value of data. When a confidence
interval is calculated, it then includes the true value with a probability of 95
percent.

Conflicto: the earliest genus of geese.

Conodonta: a clade of basal vertebrates.

convergent evolution: when completely unrelated animals evolve to share a

similar body shape, even though they did not descend from the same
ancestor.

Cope, Edward Drinker (1856–1897): an American vertebrate paleontologist,

zoologist, comparative anatomist, herpetologist, and ichthyologist.

coprolites: trace fossils consisting of feces, which are usually phosphatic.

Cretaceous: the youngest period of the Mesozoic (145 to 66 million years

before present).

Cymbospondylus: a genus of marine reptiles from the ichthyosaur clade.

cynodontia: a group of early mammal-like therapsids.

Daonella: a genus of free-swimming marine bivalves from the Triassic

period.

dicynodontia: a group of herbivorous synapsids.

dinosauria: the largest group of terrestrial vertebrates of the Mesozoic era.

Dinosauromorpha: a major group within the archosaurs that includes the

dinosaurs and their closest relatives.

Diplodocus: a genus from the group of long-necked dinosaurs from the

Upper Jurassic found in the Morrison Formation.

ecology: the study of living things’ relationships to each other and their
environment.

Edmontosaurus: a genus of duckbill dinosaurs.

Enantiornithes: a sister group of modern birds, that went extinct 66 million
years ago.

Eocene: a chronostratigraphic series of the Paleogene (56 to 33.9 million

years before present).

Europasaurus: an island-dwarfed sauropod from northern Germany.

fossil deposit: a site that is particularly rich in fossils.

fossil record: a documentation of all occurrences of fossils that places them

in a scientific, stratigraphic context.

fossils: the remains of living things and evidence of their activity that are at
least ten thousand years old and can attest to past life in Earth’s history.

Galloanserae: a major group of modern birds, including all fowl and geese.

Germanic Basin: a large sedimentary area that extended from England to

eastern Poland during the Permian and the Triassic and was at times
covered by a shallow sea.

ghost lineage: a gap in a species lineage from a hypothesized ancestor in a

species that has left no fossil evidence, but can still be inferred to have
existed because of its descendants.

Gondwana: a southern supercontinent that consisted of present-day Africa,

South America, Antarctica, India, and Australia.

Helveticosauridae: a family of basal archosauromorph marine reptiles from

the Middle Triassic of Switzerland.
Heyuannia: a genus from the oviraptorian group from China, of which
innumerable clutches are known.

Holtz, Thomas Richard, Jr. (*1965): an American vertebrate paleontologist

with the University of Maryland.

Ichthyosauria: a group of extinct secondary marine reptiles that were fully

adapted to life in the sea.

Iguanodon: a genus in the group of ornithopod dinosaurs.

island dwarfing: a phenomenon in evolutionary biology whereby the size of

animals in an isolated ecosystem decreases within a few generations. It
occurs especially on islands where these animals do not have to fear
predators. Other remote habitats such as caves, cut-off valleys, or
inaccessible mountainous regions can also cause this phenomenon.

iridium anomaly: a globally detectable elevated concentration of the

element iridium in sedimentary rocks deposited 66 million years ago, at the
Cretaceous–Paleogene boundary, which is evidence of an asteroid impact.

Janensch, Werner Ernst Martin (1878–1969): a German vertebrate

paleontologist, geologist, explorer, and one of the foremost dinosaur
specialists of his time, best known for his African expedition.

Jurassic: the middle period of the Mesozoic (201.3 to 145 million years
before present).

Kimmeridgian: the middle stage of the Upper Jurassic (157.3 to 152.1

million years before present).

L
Leidy, Joseph (1823–1891): an American paleontologist and professor of
anatomy with the University of Pennsylvania.

Maastrichtian: the latest stage of the Upper Cretaceous (72 to 66 million

years before present).

Maniraptora: a derived group of the Coelurosauria.

Maraapunisaurus (formerly Amphicoelias): a genus of long-necked

dinosaurs from the Upper Jurassic of North America found in the Morrison
Formation. The animal is considered a hot contender for the record of the
largest dinosaur ever found.

Marsh, Othniel Charles (1831–1899): an American vertebrate

paleontologist and the first professor of paleontology at Yale University.

Megalosauridae: a family within the group Megalosauroidea.

Megalosauroidea: a superfamily within the group of theropods.

Megalosaurus: a genus of theropods from the Middle Jurassic of England

(about 166 million years before present) that, in 1824, became the very first
scientifically described dinosaur.

melanosomes: the specialized functional systems in pigment cells that

produce the pigment melanin.

Mellisuga: an extant genus of hummingbirds and the smallest bird in the

world.

Mesozoic: the second-to-last era of Earth’s geological history, also known

as “Earth’s Middle Ages” (251.9 to 66 million years before present).

Mixosauria: a group of small ichthyosaurs that lived during the Middle

Triassic.

N
Neoaves: the group of all modern birds except the Paleognathae and the
Galloanserae.

Neognathae: the group of all modern birds except the Paleognathae.

Neovenator: a dinosaur closely related to Allosaurus and dated to the Lower

Cretaceous (130.7 to 126.3 million years before present).

Norian: the middle chronostratigraphic stage of the Upper Triassic (228 to

208.5 million years before present).

Notatesseraeraptor: a genus of small theropods from the Triassic closely

related to Dilophosaurus. The first carnivorous dinosaur described from
Switzerland.

Odontocetes: the scientific name for toothed whales.

Oligocene: the uppermost series of the Paleogene (33.9 to 23 million years

before present).

Omphalosaurus: an unusual ichthyosaur from the Triassic with crushing

dentition.

Ornithischia: the scientific name for the bird-hipped dinosaurs.

Ornithopoda: a clade of ornithischian dinosaurs that started out as small,

bipedal running grazers, and evolved into large duckbill dinosaurs.

Osborn, Henry Fairfield, Sr. (1857–1935): an American vertebrate

paleontologist, geologist, and president of the American Museum of Natural
History.

osteoderms: dermal bones that usually form a carapace and serve to protect
an animal.

Ostrom, John Harold (1928–2005): an American vertebrate paleontologist

and discoverer of the predatory dinosaur Deinonychus.
Ostromia: a genus of small animals from the group of Maniraptora.

Owen, Sir Richard (1804–1892): an English paleontologist, biologist,

comparative anatomist, and head of the British Natural History Museum in
London, who coined the term “Dinosauria.”

Oxfordian: the oldest stage of the Upper Jurassic (163.5 to 157.3 million
years before present).

pachyostosis: a secondary adaptation of the descendants of terrestrial

vertebrates, acquired through evolution, to life in water, in order to
compensate for the static buoyancy of the body and to be able to remain
underwater without effort. Of animals still living today, only manatees show
pachyostosis, which is particularly noticeable in their broad, thick, and
dense rib bones.

Pachypes: an ichnospecies of four-legged, five-toed herbivores from the

Permian.

Paleognathae: the group of all modern birds except the Neognathae and the
Galloanserae.

paleobond: an adhesive used to glue rocks, fossils, and petrified wood.

Paleogene: the oldest geochronologic period of the Cenozoic Era (66 to 23

million years before present).

Pangaea: a primitive continent that united all landmasses of the Earth in

itself in the Permian.

Permian: the youngest period of the Paleozoic (298.9 to 251.9 million years
before present).

paedomorphosis: the retention of juvenile characteristics at later life stages.

It may be achieved by accelerating sexual maturity relative to the rest of the
development or by delaying physical development relative to reproductive
capacity.
Phalarodon: an early genus of ichthyosaurs discovered in Nevada and
belonging to the mixosaur group.

phylogeny: within biological systematics, phylogeny describes the

phylogenetic development of all living organisms and the relationship of the
groups to each other.

Plateosaurus: a basal genus of Sauropodomorpha from the Upper Triassic.

pneumatization: the formation of air-filled cavities in originally dense bone

tissue. Pneumatization can occur during the course of evolution as well as
the course of individual development. Such bones are also referred to as
“air-filled bones.”

Potamornis: an extinct bird from the group of Hesperornithes, from the

latest Cretaceous.

precocial birds: newly hatched young that are sufficiently developed to

leave the nest and immediately follow their parents.

Proceratosauridae: a family of the group Tyrannosauroidea and sister group

to the Tyrannosauridae.

Rauhut, Oliver (*1969): a German vertebrate paleontologist at the

University of Munich.

Sander, Paul Martin (*1960): a German vertebrate paleontologist at the

University of Bonn.

Saurischia: the scientific name for the group of lizard-hipped dinosaurs.

Sauropoda: the group of long-necked dinosaurs.

Sauropodomorpha: one of the two groups within the saurischian dinosaurs.

scanning electron microscope: a microscope that passes an electron beam
over an object in a specific pattern. The interactions of the electrons with
the object are used to generate an image.

serration: a sawtooth-like cutting edge on teeth.

sexual dimorphism: the externally visible differences between sexually

mature male and female individuals of a species that do not relate to sexual
organs.

Shonisaurus: a genus of ichthyosaurs discovered in Nevada and among the

largest ichthyosaurs ever found.

Silesaurus: a genus of Dinosauromorpha from Poland and one of the closest

relatives of the dinosaurs.

sclerotic ring: a bony, ring-shaped structure around the eyes of birds,

dinosaurs, pterosaurs, ichthyosaurs, and iguanas.

Spinosauridae: a family from the group Spinosauroidea.

Spinosaurus: a genus of the Spinosauridae family with dorsal sails and one
of the largest carnivorous dinosaurs, which has been found in Morocco and
Egypt.

Stegosaurus: a genus of the ornithischian dinosaur group with bony plates

on the back and a spiny tail found in the Morrison Formation.

Sternberg, Charles Hazelius (1850–1943): an American fossil collector.

synapomorphy: a derived characteristic that groups species together and

distinguishes them from other species.

Synapsida: a large group of the amniotes.

synchrotron: a special type of particle accelerator.

T
Tanystropheus: a genus of basal Middle and Upper Triassic
archosauromorphs whose greatly elongated neck was at least as long as
their trunk and tail combined.

tsunamites: sedimentary rocks composed of the sediments formed during a

tsunami.

tektites: glass objects that are formed when surrounding rock melts during
an asteroid or meteorite impact and is hurled away from the impact site. The
mostly teardrop-shaped objects are several centimeters in length and consist
of modifications of quartz.

Temnospondyls: a group of terrestrial vertebrates that may be the ancestors

of modern amphibians.

tetrachromacy: a type of color vision with four different cone cells. Reptiles
are tetrachromats. It is believed that tetrachromats can see up to 100 million
colors. Tetrachromacy is an original characteristic of terrestrial vertebrates.

Tethys: a Mesozoic ocean, located between the ancient continents of

Gondwana and Laurasia, that completely closed during the Cenozoic, about
50 million years ago.

Tetrapoda: the large group of all terrestrial and secondarily aquatic

vertebrates.

Thalattoarchon: a large predatory genus of the ichthyosaur group.

Therapsida: a large group of terrestrial vertebrates that includes mammals.

Theropoda: a group of saurischian dinosaurs that includes all carnivorous

dinosaurs.

Thrinaxodon: a genus from the group of therapsids.

Tithonian: the uppermost stage of the Upper Jurassic (152.1 to 145 million
years before present).

Torvosaurus: a genus of the family Megalosauridae.

Triassic: the oldest period of the Mesozoic (251.9 to 201.3 million years
before present).

Triceratops: a genus from the group of horned dinosaurs.

trophic level: a particular level within the food chain in an ecosystem.

Tuebingosaurus: a genus of a sauropodomorph dinosaur from the Late

Triassic of Germany.

tuffs: volcanic igneous rocks.

Tyrannosauroidea: a superfamily from the group Theropoda.

Tyrannosauridae: a family from the group Tyrannosauroidea.

Tyrannosaurus: a genus of the group Tyrannosauridae.

Thyreophora: a group of ornithischian dinosaurs that includes all stegosaurs

and ankylosaurs.

Yutyrannus: a genus of the group Tyrannosauroidea.

zircon: a silicate mineral with the chemical formula Zr[SiO]4.

OceanofPDF.com
 References

Chapter 1: New Life in the Sea

Wintrich, T. et al. An enigmatic marine reptile—the actual first record of

Omphalosaurus in the Muschelkalk of the Germanic basin. Journal of
Vertebrate Paleontology (2017).

Fröbisch, N. B. et al. Macropredatory ichthyosaur from the Middle Triassic

and the origin of modern trophic networks. PNAS (2013).

Sander, P. M. et al. Early giant reveals faster evolution of large body size in
ichthyosaurs than in cetaceans. Science (2021).

Motani, R. et al. Terrestrial Origin of Viviparity in Mesozoic Marine

Reptiles Indicated by Early Triassic Embryonic Fossils. PLoS ONE
(2014).

Motani, R. et al. A basal ichthyosauriform with a short snout from the

Lower Triassic of China. Nature (2015).

Klein, N. et al. A new cymbospondylid ichthyosaur (Ichthyosauria) from the

Middle Triassic (Anisian) of the Augusta Mountains, Nevada, USA.
Journal of Systematic Palaeontology (2020).

Marchetti, L. et al. Leaving only trace fossils -the unknown visitors of

Winterswijk. Staringia (2019).

Wintrich, T. et al. A Triassic plesiosaurian skeleton and bone histology

inform on evolution of a unique body plan. Science Advances (2017).

Krahl, A. & Witzel, U. Foreflipper and hindflipper muscle reconstructions

of Cryptoclidus eurymerus in comparison to functional analogues:
 introduction of a myological mechanism for flipper twisting. PeerJ
(2021).

Chapter 2: New Life on Land

Fernandez, V. et al. Synchrotron Reveals Early Triassic Odd Couple:

Injured Amphibian and Aestivating Therapsid Share Burrow. PLoS
ONE (2013).

Qvarnström, M. et al. Tyrannosaurid-like osteophagy by a Triassic

archosaur. Scientific Reports (2019).

Qvarnström, M. et al. Beetle-bearing coprolites possibly reveal the diet of a

Late Triassic dinosauriform. Royal Society Open Science (2019).

Qvarnström, M. et al. Exceptionally preserved beetles in a Triassic coprolite

of putative dinosauriform origin. Current Biology (2021).

Nesbitt, S. J. et al. The oldest dinosaur? A Middle Triassic dinosauriform

from Tanzania. Biology Letters (2013).

Pol, D. et al. Triassic sauropodomorph dinosaurs from South America: The

origin and diversification of dinosaur dominated herbivorous faunas.
Journal of South American Earth Sciences (2021).

Griffin, C. T. et al. Africa’s oldest dinosaurs reveal early suppression of

dinosaur distribution. Nature (2022).

Zahner, M. & Brinkmann, W. A Triassic averostran-line theropod from

Switzerland and the early evolution of dinosaurs. Nature Ecology &
Evolution (2019).

Hofmann, R. & Sander, P. M. The first juvenile specimens of Plateosaurus

engelhardti from Frick, Switzerland: isolated neural arches and their
implications for developmental plasticity in a basal sauropodomorph.
PeerJ (2014).
Fernández, O. R. R. & Werneburg, I. A new massopodan sauropodomorph
from Trossingen Formation (Germany) hidden as ‘Plateosaurus’ for
100 years in the historical Tübingen collection. Vertebrate Zoology
(2022).

Olsen, P. et al. Arctic ice and the ecological rise of the dinosaurs. Science
Advances (2022).

Chapter 4: Germany’s Dinosaurs

Sander P. M. et al. Bone histology indicates insular dwarfism in a new Late

Jurassic sauropod dinosaur. Nature (2006).

Lallensack, J. et al. Dinosaur tracks from the Langenberg Quarry (Late

Jurassic, Germany) reconstructed with historical photogrammetry:
Evidence for large theropods soon after insular dwarfism.
Palaeontologia Electronica (2015).

Foth, C. & Rauhut, O. W. M. Re-evaluation of the Haarlem Archaeopteryx

and the radiation of maniraptoran theropod dinosaurs. BMC
Evolutionary Biology (2017).

Rauhut, O. W. M. et al. A non-archaeopterygid avialan theropod from the

Late Jurassic of southern Germany. eLife (2019).

Taylor, M. P. A re-evaluation of Brachiosaurus altithorax Riggs, 1903

(Dinosauria, Sauropoda) and Its Generic Separation from Giraffatitan
brancai (Janensch 1914). Journal of Vertebrate Paleontology (2009).

D’Emic, M. D. et al. Anatomy, systematics, paleoenvironment, growth, and

age of the sauropod dinosaur Sonorasaurus thompsoni from the
Cretaceous of Arizona, USA. Journal of Paleontology (2016).

Mannion, P. D. et al. The earliest known titanosauriform sauropod dinosaur

and the evolution of Brachiosauridae. PeerJ (2017).
 Chapter 5: Argentina—Where the Giants Live

Carballido, J. L. et al. A new giant titanosaur sheds light on body mass

evolution among sauropod dinosaurs. Proceedings of the Royal
Society B: Biological Sciences (2017).

Rauhut, O. W. M. & Pol, D. Probable basal allosauroid from the early

Middle Jurassic Cañadón Asfalto Formation of Argentina highlights
phylogenetic uncertainty in tetanuran theropod dinosaurs. Scientific
Reports (2019).

Chapter 6: Myanmar—Trapped in “Liquid Gold”

Xing, L. et al. A mid-Cretaceous enantiornithine (Aves) hatchling preserved

in Burmese amber with unusual plumage. Gondwana Research (2017).

Peñalver, E. et al. Ticks parasitised feathered dinosaurs as revealed by

Cretaceous amber assemblages. Nature Communications (2017).

Xing, L. et al. Hummingbird-sized dinosaur from the Cretaceous period of

Myanmar. Nature (2020). ARTICLE WITHDRAWN

Bolet, A. et al. Unusual morphology in the mid-Cretaceous lizard

Oculudentavis. Current Biology (2021).

Dunne, E. M. et al. Ethics, law, and politics in paleontological research: The

case of Myanmar amber. Communications Biology (2022).

Chapter 7: The Spinosaurs

Smith, J. B. et al. A Giant Sauropod Dinosaur from an Upper Cretaceous

Mangrove Deposit in Egypt. Science (2001).

Ibrahim, N. et al. Semiaquatic adaptations in a giant predatory dinosaur.

Science (2014).
Henderson, D. M. A buoyancy, balance and stability challenge to the
hypothesis of a semi-aquatic Spinosaurus Stromer, 1915 (Dinosauria:
Theropoda). PeerJ (2018).

Ibrahim, N. et al. Tail-propelled aquatic locomotion in a theropod dinosaur.

Nature (2020).

Hone, D. & Holtz, T. Evaluating the ecology of Spinosaurus: shoreline

generalist or aquatic pursuit specialist? Palaeontologia Electronica
(2021).

Sereno, P. C. et al. Spinosaurus is not an aquatic dinosaur. eLife (2022).

Barker, C. T. et al. New spinosaurids from the Wessex Formation (Early

Cretaceous, UK) and the European origins of Spinosauridae. Scientific
Reports (2021).

Barker, C. T. et al. A European giant: a large spinosaurid (Dinosauria:

Theropoda) from the Vectis Formation (Wealden Group, Early
Cretaceous), UK. PeerJ (2022).

Chapter 8: More than Just Bones

Wang, X. et al. Archaeorhynchus preserving significant soft tissue including

probable fossilized lungs. PNAS (2018).

Bell, P. R. et al. The exquisitely preserved integument of Psittacosaurus and

the scaly skin of ceratopsian dinosaurs. Communications Biology
(2022).

Vinther, J., Nicholls, R. & Kelly, D. A. A cloacal opening in a non-avian

dinosaur. Current Biology (2021).

O’Connor, J. K. et al. Ovarian follicles shed new light on dinosaur

reproduction during the transition toward birds. National Science
Review (2014).
Bailleul, A. M. et al. Confirmation of ovarian follicles in an enantiornithine
(Aves) from the Jehol biota using soft tissue analyses.
Communications Biology (2020).

Brown, C. M. et al. An Exceptionally Preserved Three-Dimensional

Armored Dinosaur Reveals Insights into Coloration and Cretaceous
Predator-Prey Dynamics. Current Biology (2017).

Chapter 9: Wyoming—The Hell Creek

Schmitt, A. D. Auf der Dinosaurier-Baustelle: Zwischen Knochen und

Kettensägen. National Geographic (August 2019). Written in German.

Schmitt, A. D. Wie verschifft man ein Stück Dinosaurier-Friedhof? National

Geographic (August 2019). Written in German.

Chapter 10: Tyrannosaurus—The Measure of All Things!

Marshall, C. R. et al. Absolute abundance and preservation rate of

Tyrannosaurus rex. Science (2021).

Carpenter, K. & Smith, M. Forelimb Osteology and Biomechanics of

Tyrannosaurus rex. Mesozoic vertebrate life. Bloomington: Indiana
University Press (2001).

Carbone, C. et al. Intra-guild competition and its implications for one of the
biggest terrestrial predators, Tyrannosaurus rex. Proceedings of the
Royal Society B: Biological Sciences (2011).

Holtz, T. R. Theropod guild structure and the tyrannosaurid niche

assimilation hypothesis: implications for predatory dinosaur
macroecology and ontogeny in later Late Cretaceous Asiamerica.
Canadian Journal of Earth Sciences (2021).

Schroeder, K., et al. The influence of juvenile dinosaurs on community

structure and diversity. Science (2021).
Carr, T. D. A high-resolution growth series of Tyrannosaurus rex obtained
from multiple lines of evidence. PeerJ (2020).

Bell, P. R. et al. Tyrannosauroid integument reveals conflicting patterns of

gigantism and feather evolution. Biology Letters (2017).

Padian, K. Why tyrannosaur forelimbs were so short: An integrative

hypothesis. Acta Palaeontologica Polonica (2022).

Bhullar, B.-A. S. et al. Birds have paedomorphic dinosaur skulls. Nature

(2012).

Ullmann, P. V. et al. Taphonomic and Diagenetic Pathways to Protein

Preservation, Part I: The Case of Tyrannosaurus rex Specimen. MOR
1125. Biology (2021).

Paul, G. S. et al. The Tyrant Lizard King, Queen and Emperor: Multiple
Lines of Morphological and Stratigraphic Evidence Support Subtle
Evolution and Probable Speciation Within the North American Genus
Tyrannosaurus. Evolutionary Biology (2022).

Carr, T. D. et al. Insufficient Evidence for Multiple Species of

Tyrannosaurus in the Latest Cretaceous of North America: A Comment
on “The Tyrant Lizard King, Queen and Emperor: Multiple Lines of
Morphological and Stratigraphic Evidence Support Subtle Evolution
and Probable Speciation Within the North American Genus
Tyrannosaurus.” Evolutionary Biology (2022).

Chapter 11: Movement Captured in the Stone: What Footprints Tell Us

Schanz, T. et al. Quantitative Interpretation of Tracks for Determination of

Body Mass. PLoS ONE (2013).

Lockley, M. et al. Titanosaurid trackways from the Upper Cretaceous of

Bolivia: Evidence for large manus, wide-gauge locomotion and
gregarious behaviour. Cretaceous Research (2002).
 Chapter 12: Birds—The Last Dinosaurs

Prum, R. O. et al. A comprehensive phylogeny of birds (Aves) using

targeted next-generation DNA sequencing. Nature (2015).

Cloutier, A. et al. Whole-Genome Analyses Resolve the Phylogeny of

Flightless Birds (Palaeognathae) in the Presence of an Empirical
Anomaly Zone. Systematic Biology (2019).

Wiemann, J., et al. Dinosaur egg colour had a single evolutionary origin.
Nature (2018).

Field, D. J. et al. Late Cretaceous neornithine from Europe illuminates the

origins of crown birds. Nature (2020).

Koschowitz, M.-C. et al. Beyond the rainbow. Science (2014).

Foth, C. et al. New specimen of Archaeopteryx provides insights into the

evolution of pennaceous feathers. Nature (2014).

Smithwick, F. M. et al. Countershading and Stripes in the Theropod

Dinosaur Sinosauropteryx Reveal Heterogeneous Habitats in the Early
Cretaceous Jehol Biota. Current Biology (2017).

Hu, D. et al. A bony-crested Jurassic dinosaur with evidence of iridescent

plumage highlights complexity in early paravian evolution. Nature
Communications (2018).

Chapter 13: The End of the Dinosaurs

During, M. A. D. et al. The Mesozoic terminated in boreal spring. Nature

(2022).

Alvarez, L. W. et al. Extraterrestrial Cause for the Cretaceous-Tertiary

Extinction. Science (1980).
Santa Catharina, A. et al. Timing and causes of forest fire at the K–Pg
boundary. Scientific Reports (2022).

Nicholson, U. et al. The Nadir Crater offshore West Africa: A candidate

Cretaceous-Paleogene impact structure. Science Advances (2022).

Field, D. J. et al. Early Evolution of Modern Birds Structured by Global

Forest Collapse at the End-Cretaceous Mass Extinction. Current
Biology (2018).

OceanofPDF.com
 Acknowledgments

When I was in seventh or eighth grade, my English skills were below

average, and my English teacher told me that I would never learn how to
speak proper English, let alone write a single concise sentence. Yet here I
am. It is virtually impossible for other people to know what we are capable
of until we do it. We all have skills and talents in us that are not apparent to
others, and sometimes it just needs a little push and some encouragement
from friends, family, and mentors for us to show the world what we are
capable of. They may not know if we succeed, but they have faith in us and
trust in our potential. That is why I would like to thank those who made it
possible for me to write this book. This is most certainly not a complete list
—there may be a few people whom I forgot to mention, and I apologize for
that in advance.
I am particularly grateful for all the help and support I received from
Martin Sander in Bonn, who was my supervisor for my diploma thesis. He
recognized my potential and gave me a chance to prove myself, but I also
learned so much from him that I am now myself able to pass on my
knowledge to others. Equally important for my career in paleontology is
Roger Benson, whom I worked for as a research assistant in Oxford. I can’t
overstate how lucky I feel to have had the opportunity to work with one of
the most brilliant minds I have ever encountered in my life. If it wasn’t for
this job, many other consecutive career steps would not have been possible.
I owe gratitude to Daniel Field for offering me a doctoral research position
in Cambridge. Had it not been for this position, I would have never been
interviewed by Frank Thadeusz from the news journal Der Spiegel. Thanks
to him, people started taking an interest in my dinosaur content on various
social media platforms, and so it was him who pointed the publisher DTV
in my direction. Here, I would like to thank Laura Weber, who offered me
the opportunity to write a book about my favorite topic: dinosaurs. And
Andrea Seibert, who did a tremendous job at presenting my popular science
book to a list of international publishers at the London Book Fair. I am very
thankful to Hanover Square Press for giving me the opportunity to present
my book to an English-speaking audience and thus sharing my love for
dinosaurs with people across the globe. At Hanover Square Press, Eden
Railsback and Peter Joseph did a phenomenal job with helping me convert
this book into a presentable English version. They both have been
supportive and incredibly encouraging from the moment we started working
together.
And while the book is out now and I am proud of my accomplishment,
the book came about in a time when I was struggling with severe personal
problems, and had it not been for friends like Sabrina Müller and the
wonderful Maryness Honoratus, I would not have been able to push through
and overcome all these challenges. I would also like to thank Ben Marshall
from Hughes Hall College for providing mental support, and Philip Isaac
from King’s College, who helped get me back on my feet. In the end, it is
always the success that other people see, and never the struggles we
encounter along the way.
A special thanks goes out to Steve Brusatte, who has always encouraged
me to continue to pursue my career in paleontology and to write a book, and
to Thomas Holtz, who has been very supportive during the writing process
of this book by providing valuable information and insight into the anatomy
and physiology of Tyrannosaurus. I would also like to thank my former
housemate Davinder, who helped me with an earlier translation of two
chapters of this book, albeit his translation ultimately did not end up in the
final product. Nevertheless, I acknowledge and appreciate his effort. Of
course, I would like to thank Lars Schmitz and Georg Oleschinski for
letting me use their photos for this book!
Lastly, I would like to thank my family for making me the person I am
today. I owe everything to my mother and my father, who may not have
always understood me but who always made sure I felt loved
unconditionally and always accepted me for who I am, despite being
difficult or awkward at times. I would also like to thank my brother,
Michael, and my sister, Sonja, for keeping up with me, despite being the
most annoying youngest sibling imaginable.
Most importantly, all my love goes out to my beautiful children,
Maximilian and Elisabeth, who are the most precious and wonderful human
beings imaginable. They are in my heart always and they are the apples of
my eye and the reason and motivation behind my achievements. I hope they
feel loved unconditionally by their dad, as I feel loved by my parents.
OceanofPDF.com
OceanofPDF.com
OceanofPDF.com
 Index

The pagination of this digital edition does not match the print edition from
which the index was created. To locate a specific entry, please use your
ebook reader’s search tools.

A
abelisaurians, 218–19
Abelisaurus, 218–19
Aegyptosaurus, 147, 148
aestivation
described, 61
growth and, 64
torpor stage compared to, 65
aetosaurs, 52
Aiglstorfer, Manuela, 188, 189
Alamosaurus, 127, 203
Albertosaurus, 214–15
Alcmonavis, 109
Allosaurus, Europasaurus and relatives of, 105
Alvarez, Luis, 248
Alvarez, Walter, 248
amber
from Canada, 132
fossils in, 131–33, 137, 138, 140–41
from Myanmar, 132, 140, 142–44
American Museum of Natural History (New York City), 82
ammonites, 26(illus)
basic facts about, 17
oxygen-deficient environments and, 35
as prey of Omphalosaurus, 31
recovery of, after “the Great Dying,” 43
amniotes, 59–60
Amphicoelias fragillimus, 96
anagenesis, 223
anatomy
abelisaurians, 218–19
ankylosaurs, 161
Archaeopteryx, 107(illus), 247
Archaeorhynchus, 165–66
archosaurs, 67–68, 167–69
Baryonyx, 163
birds, 168–69, 219
Brachiosaurus, 110
differentiating into species based on, 221–24
dinosaurs, 68–69, 138, 242
Diplodocus, 232
Edmontosaurus, 211
Hypsilophodon, 161
ichthyosaurs, 29, 138–39
Iguanodon, 160, 161
lack of current animals with comparable, 14
learned from examining bones, 227
lungs, 165
lungs’ oxygen capacity, 21, 84
Oculudentavis, 138–39, 140
ornithischian dinosaurs, 73
paleognath birds, 238
Plateosaurus, 76
sauropods, 215
“secondary metamorphosis,” 210–11
Silesaurus, 73, 181
Spinosaurus, 151, 152–54, 155, 156–58
theropods, 219
Triceratops, 178–79, 180–81
Tyrannosaurus, 210–11, 212
Tyrannosaurus rex, 217–18
See also dentition
ankylosaurs
anatomy, 161
Borealopelta markmitchelli, 170–173, 227
Polacanthus, 161
tracks in south America, 233
Ankylosaurus, 203
Anning, Mary, 28
apex predators
described, 40
in ecosystems, 40, 173, 199
eyesight, 173–74
aposematism, 173
Archaeopteryx
anatomy, 107(illus), 247
feathers of, 109, 247–48
first scientific description of, 94, 106
Archaeorhynchus, 165–66
archosaurs
anatomy, 67–68, 167–69
dentition of, 66
Dinosauromorpha, 71, 72–73
“the Great Dying” and, 21
locomotion, 67
split into crocodiles and dinosaurs, 66–67, 240
synapomorphies of, 66
Argentinosaurus, 119, 122, 123
anatomy, 110
discovery of, 112–13, 119
phylogenetic tree of, 113–15
size of, 102, 119
arrested growth, 64
Asaro, Frank, 248
Asteriornis, 261
Asteriornis maastrichtensis (“Asteria’s bird”), 245
Augusta Mountains (Nevada), 27–28, 30, 47, 48–50
autochthonous fossils, 228
B
Bahariasaurus, 147
Bahariya Basin (Egypt), 146–48, 163–64
Bakker, Robert, 109, 195
Barnes High sauropod, 161
Baryonyx, 118(illus), 162–63
bee hummingbirds (Mellisuga helenae), 121
beer, 38–39
benthos, 35
Benton, Michael, 18
biostratigraphy, 43
bioturbation, 35
bipedalism
advantages of, 83
dinosaurs and, 73
Hypsilophodon, 161
of Plateosaurus, 76
birds
Alcmonavis and, 109
anatomy, 168–69, 219
Archaeorhynchus and, 165
Chicxulub survival impact by, 260
crown birds, 244–46
current decline of, 23–24, 266
as descendants of dinosaurs, 108
eggs of, 134
enantiornithines and, 132–35, 137–38
evolution, 219–20
lungs of, 166
Marsh and relationship of, to dinosaurs, 94
mousebirds, 260–61
Oculudentavis and, 138–41
oldest fossil, 109–10
passerines, 260–61
reproduction, 134, 168–69, 241–42
Sander prediction about proto-, from Langenberg quarry, 109
 sexual characteristics of, 220
See also Archaeopteryx; neognath (“new jaw”) birds; paleognath (“old
jaw”) birds
Bismarck, Otto von, 110
Bison alticornis, 179
bivalves, 26(illus)
Black Hills Institute (South Dakota), 195, 197
black shales, 35
bone beds, removal of, 188–89, 189(illus), 190–92
Bone Wars, 91–93, 95, 97
Borealopelta markmitchelli, 170–73, 227
brachiosaurids, 104
See also Europasaurus
brachiosaurs, 232
Brachiosaurus
anatomy, 110
discovery of, 112–13, 119
phylogenetic tree of, 113–15
size of, 102, 119
broad-gauge trackways, 232
Brontosaurus, 121
Broomistega, 56(illus), 61–64
Brown, Barnum, 186, 194
Buckland, William, 92
Bull, John, 146
Burma. See Myanmar
Burpee Museum of Natural History (Illinois), 221
burrowing lifestyle
benefits of, 58, 64–65
in Karoo Basin, 58–59
Thrinaxodon, 61
Thrinaxodon and Broomistega together, 61–64

C
Caihong juji (“rainbow dinosaur”), 253–54
Cal Orck’o mountain paleontological site (Bolivia), 233
Camarasaurus, 102
camouflage, 172–74, 241
Canada, 132
Cañadón Calcáreo Formation, 126, 127
Carballido, José, 122, 123
Carbone, Chris, 201, 205, 206
Carcharodontosaurus saharicus, 147, 148
Carnotaurus, 218–19
Carpenter, Kenneth, 201
Carrier’s constraint, 67
cassowaries, 235, 238, 241, 243, 251
cephalopods and “the Great Dying,” 43
ceratopsians, 184
Ceratosuchops, 163
Cetiosaurus, 121, 161
Chicxulub impact
birds and, 260
described, 255–56
earthquakes and, 256, 257–59
ecological collapse from, 259–60, 262
fish and, 256–57, 261–62
chirotheria, 52
Chubut Province, Argentina, 127(illus), 128
cladogenesis, 223
Claosaurus annectens, 182
Clearwater Formation (Canada), 170, 173
climate change, 19, 20, 24
cloacas, 166–68
Cody, William, 97 (aka Buffalo Bill)
Coelurosaur (hollow-tailed lizards), 214, 216
Como Bluff site (Wyoming), 96
compsognathids, 251–53
Compsognathus, 169
concretions, 31, 51
Conflicto antarcticus, 261
Confuciusornis, 253
conodonts, extinction of marine, 21, 84
contour feathers, 216, 246, 247–48, 249, 250–51, 253
convergent evolution of marine animals, 29
Cope, Edward Drinker
basic facts about, 93, 94, 136
competition with Marsh of, 91, 92–93, 94–95, 97, 136
Elasmosaurus and, 95
Native Americans and, 184
Osborn and, 96
Triceratops and, 194
Tyrannosaurus rex bones and, 194
coprolites, 70, 71–72
coral reefs, 34–35
countershading, 172–74
Crichton, Michael, 131–32
crocodiles and archosaurs and, 66–67, 240
crocodilian lineage and end-Triassic mass extinction, 21–22
crown birds, 244–46
Cymbospondylus
species of, from Augusta Mountains, 47
success of, in Middle Triassic, 51
Cymbospondylus duelferi, 51
Cymbospondylus youngorum, 26(illus)
as apex predator, 45
diet of, 45
discovery of, 40
preservation in field of, 44
removal of skeleton of, from discovery site, 44
size of, 44–45
cynodonts. See Thrinaxodon

D
Dakotaraptor, 210
“the dance floor,” 52
daonellids, 35
Darwin, Charles, 93, 106
Daspletosaurus, 214–15
decomposers, 41
deformation of bones, 141
Deinonychus, 108
dentition
archosaurs, 66
differentiating into species based on, 224
dinosaurs, 69
environmental information from, 188
Neovenator, 162
Oculudentavis, 139, 141
Omphalosaurus, 32, 33, 36
placodonts, 34, 36, 53
Silesaurus, 72
Spinosaurus, 149, 157
Thalattoarchon saurophagis, 40
Tyrannosaurus, 69, 149
Tyrannosaurus rex, 194
Denversaurus, 176(illus)
Dicraeosaurus, 112
Dinornis, 237
Dinosaur 13 (movie), 197
“dinosauria,” term coined, 92
Dinosaur Island of England (Isle of Wight), 159(illus), 159–64, 162(illus)
Dinosauromorpha, 71, 72–73
Dinosaur Park Münchehagen (Germany), 161
the “dinosaur plateau,” 228–29
dinosaurs
age of, and diet differences, 262
anatomy, 68–69, 138, 242
archosaurs and, 66–67, 240
behavior of, 228
bipedalism and, 73
birds as descendants of, 108
camouflage, 172
determining body mass of, 122
 eggs of, 242–43
end-Triassic mass extinction of competitors of, 84
evolution of, 74, 103–4
eyesight, 249, 253
feathers, 253
feeding behavior of, 69
first American found, 92
first excavations of, 82
footprints on Isle of Wight, 159(illus), 160
“the Great Dying” and, 20
length of time of reign of, 22
Marsh and relationship of, to birds, 94
medium-sized carnivores during Late Upper Cretaceous of North
America, 203
mortality and number of eggs laid, 203
numbers of carnivores compared to herbivores, 207
ontogenetic development of, 103–4
origin of, 73–74
plumage development by, 84
protofeathers, 248
reproduction, 203, 208, 242–43, 248
rethinking of biology of, 108–9
scales, 248
sexual characteristics, 220
term coined, 77
therapsids and, 60
tracks, 160, 161
See also specific dinosaurs
Dinosaur Valley State Park (Texas), 228–29
Diplodocus, 232
Drawno Beds Formation, 181
Drevermann, Fritz, 185
Dülfer, Olaf, 51, 104
During, Melanie, 257
dwarfism, 102–3, 138, 139, 220
E
earthquakes and Chicxulub impact, 256, 257–59
ecosystems
apex predators in, 40, 173, 199
Chicxulub impact and, 259–60, 262
coexistence of different genera in, 210
current destruction of bird, 266
expedition to understand, of Edmontosaurus, 186–92
extinctions and collapse of, 266
feathers and, 215
“the Great dying” and, 18
island dwarfing and, 103
“niche assimilation” in, 211
predators in Tyrannosaurus-dominated, 205, 206–7
preservation of fossils in forest, 140
trophic levels in, 41
Edmontosaurus, 176(illus), 183(illus)
anatomy, 211
expedition to understand ecosystem of, 186–92
first discovery of skeleton of, 184
in Hell Creek and Lance formations, 203
size of, 185, 211
Trachodon mummy, 181–82
Elasmosaurus, 95
elephant birds, 235, 236, 237
El Fayum (Egypt), 145
emus, 235, 238, 243
enantiornithines, 132–35, 137–38, 168, 169, 262
end-Cretaceous mass extinction, 17, 22
See also Chicxulub impact
end-Triassic mass extinction, 21–22, 84
Engelhardt, Friedrich, 76
Europasaurus, 100(illus)
distribution of, 104
dwarfing of, 102, 220
extinction of, 105
 first discovery of, 101
Sander and, 102
Schmitt and, 100–101, 104
size of, 102
study of ontogenetic stages of, 103–4
European Synchrotron Radiation Facility (ESRF), 57, 58, 70
evolution
of birds, 219–20, 238–39
continental drift and, 103–4
convergent, of marine animals, 29
of dinosaurs, 74, 103–4
of ichthyosaurs, 28–29, 45, 51–52
publication of theory of, 93, 106
of sauropods, 126–27
extinctions
causes of, 17, 18–20, 21
ecological collapse and, 266
end-Cretaceous mass, 17, 22, 259 (see also Chicxulub impact)
end-Triassic mass extinction, 21–22, 84
energy requirements and, 261, 262
of Europasaurus, 105
predator sizes and, 203–4, 205–6
of Stegosaurus, 22–23
“the Great Dying,” 18–20, 21
eyesight
apex predators, 173–74
color spectrum, 249, 250
cone cells, 249
dinosaurs, 249, 253
ostrich, 236
paleognath birds, 251

F
feathers
advantages of, 85
of Archaeopteryx, 109, 247–48
 Bavarian “Urvögel,” 72
body mass and, 215
Caihong juji, 253–54
coloration of, 248–49
development of, 84
dinosaurs, 253
ecosystems and, 215
enantiornithines, 135
filamentous, 133, 248, 251
Maniraptora, 249–51
metabolic rate and miniaturization and, 248, 250
microraptors, 253
oldest evidence in fossil record of, 72
ostriches, 251
paleognath chicks, 236
pennaceous (contour), 216, 246, 247–48, 249, 250–51, 253
planar, 250
precocial and altricial characteristics, 133–34
preserved in amber, 132, 133
protofeathers, 248
reproduction and, 249
Sinosauropteryx, 252
ticks on, 135
transformation of, into scales, 216
Tyrannosauroidea, 214
Tyrannosaurus rex, 213, 216
von Meyer and, 106
Yutyrannus, 213–14, 215
Fernandez, Vincent, 57, 58, 61
Field, Daniel, 245
Field Museum (Chicago), 124, 195, 220
filamentous feathers, 133, 248, 251
first-order consumers, 41
fish and Chicxulub impact, 256–57, 261–62
“fish lizards,” 28
See ichthyosaurs
food chain, 40–41
fossil record
amber and, 131–33, 137, 138, 140–41, 142
of birds, 260
deformation and retrodeformation of bones, 141
described, 22
forest creatures and, 140
gap in paleognath birds’, 239
information from autochthonous fossils, 228
method to determine if object is bone, 129
missing links in, 106
paleognath birds, 244
trace fossils, 59, 228
Foth, Christian, 247–48
Fraas, Eberhard, 111–12
Frick site, Germany, 76, 77–78, 78(illus), 79, 80–81

G
galliformes, 236, 244, 245, 260, 261
Galloanserae, 236, 244–46
galloping greenhouse effect, 20
geobionts, 35
German East Africa, 110–11
Germanic Basin, 33–34, 52–53
Germany, 36–37, 99, 110–11
gigantism, 45, 215
Gilmoremys, 176(illus)
Giraffatitan brancai, establishment of genus, 114
global warming, 19, 20, 24
goats, dwarf, 103
Gondwana, 239–40
Gorgosaurus, 214–15, 217
Great Basin Brewing Co., 39, 45
Great Basin Collared Lizards, 30–31
“the Great Dying”
cephalopods and, 43
 coral reefs and, 34
described, 19–20
dinosaurs and, 20
ecosystems and, 18–19, 20
recovery of ammonites after, 43
growth interruption, 64

H
habitat adaptation, 106
Hatcher, John Bell, 184
Havlik, Philipe, 188, 189, 191, 192
Heilmann, Gerhard, 108
Hell Creek Formation, 177, 178, 184, 194, 195, 203, 210, 256
helveticosaurs, 53
Henderson, Donald, 153–54, 155
Hendrickson, Sue, 195
Hesperornis, 108
Heyuannia, 243, 244
hippos, dwarf, 103
Holtz, Thomas, 156, 157
Homo floresiensis, 103
Hone, David, 156, 157
Huincul Formation (Argentina), 206
humans, emergence of, 22
Huxley, Thomas Henry, 108
Hypsilophodon, 161

I
Ibrahim, Nizar, 150–52, 155, 157
ichnofossils, 59
Ichthyornis, 108
ichthyornithids, 261–62
Ichthyosaur India Pale Ale (Icky), 39
ichthyosaurs, 26(illus)
anatomy, 29, 138
countershading as survival strategy, 172
 evolution of, 28–29, 45, 51–52
first complete skeleton found, 28
locomotion, 29
nickname, 28
ocean oxygen levels and, 36
origin of, 29–30
Sander and, 34
skeleton, 29
viviparous, 47, 49–50
See also specific ichthyosaurs
Iguanodon, 118(illus), 160, 161
insects, current decline of flying, 24
“integument,” 213
International Code of Zoological Nomenclature (ICZN), 74, 114
inverse tree line, 27
iridium anomaly, 258–59
island dwarfing
described, 102–3
of Europasaurus, 102, 220
of Oculudentavis, 138, 139
Isle of Wight, 159(illus), 159–64, 162(illus)

J
Jäger, Kai, 169–70
Jane (Tyrannosaurus rex), 221
Janensch, Werner, 110, 111, 112–13, 119
Janenschia, 111
Jehol bioprovince (China), 168–69, 252–53
Jeholornis, 168–69
Jiufotang Formation (China), 165
Johnson, Dwayne “The Rock,” 197–98
Jurassic Park (Crichton), 131–32
Jurassic Park (movie), 193

K
Karoo Basin (South Africa), 58–59
Kem Kem Basin (Morocco), 150
kiwis, 235, 236
Kori bustard, 238
Kruger National Park (South Africa), 207
Kugitangtau Mountains (Turkmenistan), 228–29

L
LAGs, 64
Lance Formation, 177, 179, 184, 186, 187, 203
Langenberg quarry (Harz Mountains, Germany), 101–2, 103, 105, 109
Larson, Pete, 197
Leidy, Joseph, 92
Lessemsaurus, 82
Lipoid Foundation, 186
Lithornis celetius, 239, 240
locomotion
Archaeopteryx, 247–48
archosaurs, 67
brachiosaurs, 232
Diplodocus, 232
flight and weight, 238
ichthyosaurs, 29
Maniraptora and, 242
paleognath birds, 235
Plateosaurus, 76
sauropods, 231, 231(illus)
Silesaurus, 72–73
Spinosaurus, 157
thunniform, 29
titanosaurs, 232
Lommiswil (Switzerland), 132, 230–31, 231(illus), 232
London Museum of Natural History, 184, 185
López de Bertodano Formation, 261
Lourinhã Formation (Portugal), 91
Lower Triassic
biostratigraphic classification in, 43
 conditions on land during, 57
Mustang Canyon of Augusta Mountains, 30, 48–50
Lüdtke, Holger, 101
lungs
bidirectional compared to unidirectional, 68–69, 165
birds, 166
oxygen capacity of, 21, 84
Lusk, Wyoming, 186–87
Lystrosaurus Assemblage Zone (Karoo Basin), 58

M
Majungasaurus, 218–19
mammals, 262
mammoths, dwarf, 103
Maniraptora, 242, 243, 244, 249–51
Manospondylus gigas, 194
Maraapunisaurus, 96
marine reptiles, extinction of, 17
Markgraf, Richard, 147
Marsh, Othniel Charles
basic facts about, 93, 94, 136
birds as descendants of dinosaurs, 108
Edmontosaurus annectens and, 182
competition with Cope of, 91, 92–93, 94–95, 97, 136
Native Americans and, 184
relationship of dinosaurs to birds and, 94
Triceratops and, 179
Martina, 47
Mbiresaurus, 75
medullary bone tissue, 220–21, 227
medullary cavity, 151, 158
mega-volcanism in Russia and “the Great Dying,” 18–19
melanin, 171
Mesozoic, Germany during, 99
metabolism, plasticity in, 65
Meyer, Hans, 111
Michel, Helen, 248
microraptors, 253
Middle Triassic, 30, 40
Mitchell, Mark, 171
moas, 235, 237
Monte Carlo simulation, 199–200
Monte San Girogio, 53
moraines, 84, 85
Mörnsheim Formation, 109
Morrison Formation (United States Midwest), 91, 97, 205
Motani, Ryosuke, 33, 34, 49–50
mousebirds, 260–61
Münchehagen, dinosaur tracks at, 230
Museum of Natural History (Abu Dhabi), 198
Myanmar, 132, 140, 142–44

N
nandus, 235, 238, 241
Nanotryrannus lancensis, 195, 196(illus)
Nanuqsaurus, 214
narrow-gauge trackways, 232
National Geographic, 151
National Geographic Germany, 186
National Museum of Natural History (Washington, DC), 223
Natural History Museum (Berlin), 107, 195
Natural History Museum (London), 107, 107(illus), 194
Natural History Museum (Stuttgart), 82
natural selection, 106
Nature (journal), 137, 139, 155, 257
Nature and Biodiversity Conservation Union (NABU), 23–24
Nautilus, 17, 26(illus)
necks
Amphicoelias fragillimus, 96
Cope and Elasmosaurus reconstruction, 95
Great Basin Collared Lizards, 31
ichthyosaurs, 29
 Plateosauru, 76
sauropods, 120
Tanystropheus, 53
Neoaves, 236, 260–61
neognath (“new jaw”) birds, 235, 236, 246
Neovenator, 105, 118(illus), 162
“niche assimilation,” 211, 212
nicknames of dinosaurs
“Big John,” 178 (see also Triceratops)
“fish lizards,” 28
“Sleeping Beauty,” 172
“Swabian lindworm,” 77
Trachodon mummy, 181–82 (see also Edmontosaurus)
nomenclature, 74, 114–15
Notatesseraeraptor, 80
nothosaurs, 53
Nyasasaurus, 73–74

O
oceans during “the Great dying,” 20
Oculudentavis, 138–41
Omphalosaurus, 26(illus)
dentition, 32, 33, 36
diet, 31, 33, 34, 39
preparation of skeleton, 51
preservation in field of, 32–33
removal of skeleton of, from discovery site, 37, 38(illus)
Sander and, 34
size of, 31
On the Origin of Species (Darwin), 106
The Origin of Birds (Heilmann), 108
ornithischian dinosaurs
anatomy, 73
development of, 181
Hypsilophodon, 161
Iguanodon, 118(illus), 160, 161
 Psittacosaurus, 166–68, 167(illus)
Ornithopsis, 161
Osborn, Henry Fairfield
arms of Tyrannosaurus rex, 218
Cope and, 96
dinosaurs named by, 194
Tyrannosaurus rex reconstruction by, 217
osteoderms, 161
osteophagy, 69, 70
ostriches
aggressiveness of, 238
eggs of, 236
eyesight, 236
feathers, 251
ghost lineage among relatives, 260
size of, 237
Ostrom, John, 108–9
Ostromia, 108
ovaries, 168
oviraptors, 243
Owen, Sir Richard, 77, 92
oxygen levels
current, 18
during Late Triassic, 67
during Permian, 18
upright posture and, 67, 84

P
Pachypes dolomiticus, 47–48
pachypleurosaurs, 53
Padian, Kevin, 218
paedomorphosis, 103, 139, 220
paleognath (“old jaw”) birds
anatomy, 238
color vision of, 251
diversity of, 239, 240
 feathers of chicks, 236
in fossil record, 244
neognath birds compared to, 235
origin of, 238–39
reproduction, 236
species of, 235
Pandoravenator, 129
Pangaea, 57
Paralititan stromeri, 148
passerines, 260–61
Patagotitan, 123–24
Paul, Gregory S., 113
Peabody, George, 93
peer review process, 136
pennaceous feathers, 126, 246, 247–48, 249, 250–51, 253
Permian, Earth during, 18
Permian–Triassic mass extinction. See “the Great Dying”
Petey (Tyrannosaurus rex), 221
Phalarodon, 46, 46(illus)
placodonts, 34, 36, 53
planar feathers, 250
“plateosaurs,” 82–83
Plateosaurus, 56(illus), 77(illus), 78(illus)
basic facts about, 76
developmental plasticity of growth of, 81
diet of, 78
first remains of, found, 76–77
global discoveries of, 82
juveniles, 80–81
nickname of, 77
as prey, 79
sites in Europe with, 76, 77–78, 79, 80–81
playa strata, 79
plesiosaurs, 53–55, 54(illus)
plumage. See feathers
poachers, 49
Polacanthus, 161
Potamornis, 176(illus)
primary producers, 41
Prince Creek Formation (Alaska), 214
Procolophonichnium, 53
prosauropods, 83–84
protofeathers, 248
Psittacosaurus, 166–68, 167(illus)

Q
Qvarnström, Martin, 70, 71–72

R
Rauhut, Oliver, 109, 124–26, 128–30
rebbachisaurids, 96
Red Nose Point (Augusta Mountains), 28
regurgialites, 70
reproduction
birds, 134, 241–42
dinosaurs, 203, 208, 242–43, 248
egg coloration, 241–42, 243–44
enantiornithines, 168, 169
feathers and, 249
Jeholornis, 168–69
limit to egg size, 237–38
medullary bone tissue and, 220–21
paleognath birds, 236–37
Triceratops, 180
research, funding and publishing, 136–37
retrodeformation of bones, 141
Rhaeticosaurus, 54
rhinoceros, dwarf, 103
rhynchosaurs, 52
Riggs, Elmer, 113
Riojasaurus, 82
Riparovenator, 163
Royal Tyrrell Museum (Drumheller, Canada), 170

S
Sacrison, Stan, 195
Sander, Martin, 38(illus)
affinity of Omphalosaurus to ichthyosaurs, 34
basic facts about, 32
Europasaurus and, 102
field designations of skeleton and, 46–47
juvenile Plateosaurus and, 80–81
prediction about proto-birds from Langenberg quarry, 109
Sao Khua Formation (Thailand), 91
sauropodomorphs, basic facts about, 75
sauropods
anatomy, 215
Argentina as cradle of, 126
Barnes High, 161
Brontosaurus, 121
diet, 232
evolution of, 126–27
locomotion, 231, 231(illus)
names of, 121
necks of, 120
size of, 120–22
tracks, 228, 230–31x, 231(illus)
uniqueness of, 119–20
See also Brachiosaurus
Schmitt, Armin
Borealopelta markmitchelli and, 170, 172–73
characteristics of, 47, 48
childhood love of dinosaurs, 12, 99, 121, 177, 181, 182(illus)
dinosaurs in southern Argentina and, 124–26, 128–30
Europasaurus and, 100–101, 104
expedition to understand ecosystem of Edmontosaurus and, 186–92
field designations of skeleton and, 50
in Harz Mountains, 101–2
 Jäger and, 170
at Lommiswil, 230
reconstruction of world’s largest dinosaur by, 122–23
Spinosaurus as semiaquatic and, 151–53, 154, 155–56
Schmitz, Lars, 32, 46, 46(illus)
Schweinfurth, Georg August, 145
Science (journal), 137, 151
Science Slam, 169
sea levels, at Triassic–Jurassic boundary, 84
secondary marine animals, 151
“secondary metamorphosis” anatomy, 210–11
second-order consumers, described, 41
Seismosaurus, 121
Senckenberg Natural History Museum (Germany), 13, 177, 181, 182(illus),
185, 186
Sereno, Paul, 157–58
Siberian traps, 18–19
Silesaurus, 71, 72–73, 181
Sinosauropteryx, 251–53, 254
“Sleeping Beauty” (Borealopelta markmitchelli), 170–73, 227
Smith, Matt, 201
Smok wawelski, 70–71
Society of Vertebrate Paleontology (SVP), 142–43
Solnhofen Limestone (Bavaria), 247
“Solothurn Turtle Limestone,” 231
Sonorasaurus, 115
species, discovery of new, 14
spinosaurids, 163–64
See also specific dinosaurs
Spinosaurus
anatomy, 151, 152–54, 155, 156–58
bite force of, 150
center of mass of, 154
diet of, 150, 157
discovery of, 147–48
locomotion, 157
 as semiaquatic, 150–51, 152–58
size of, 149
squamates, 139–40
Stan (Tyrannosaurus rex), 195, 197, 198(illus)
Stegosaurus, extinction of, 22–23
Stein, Koen
basic facts about, 32
field designations of skeleton and, 46, 49
removal of Omphalosaurus skeleton, 37, 38(illus)
Sternberg, Charles Hazelius, 182–83, 184, 185, 186
Sternberg, Charles Mortam, 184–85
Sternberg, George, 184, 185
Sternberg, Levi, 184, 185
Stromer (von Reichenbach), Karl Heinrich Ernst Freiherr
Bahariya Basin discoveries by, 146–48
basic facts about, 145–46
Bull and, 146
National Socialists and, 148
Sue (Tyrannosaurus rex), 195, 197, 220, 223
Sumatran rhinoceros, 103
“Swabian lindworm,” 77
See also Plateosaurus
synapomorphies, described, 66

T
Tanis Fossil Site (North Dakota), 256
Tanystropheus, 53
Tarbosaurus, 198, 214–15
taxon, described, 74
Taylor, Mike P., 113–14
tektites, 256
temnospondyls, extinction of, 22
Tendaguru Formation (German East Africa), 111
tetrapods, described, 59
Thalattoarchon saurophagis, 26(illus)
dentition of, 40
 diet of, 40–41
as first apex predator of seas, 40
size of, 40
Thayer, Abbott, 172
Thayer’s principle, 172
“the Great Dying,” 21
therapsids
dinosaurs and, 60
LAGs in, 64
Thrinaxodon, 56(illus), 60–64, 65, 257
trace fossils of, 59
theropods, 105, 121, 173, 219
Thrinaxodon, 56(illus), 60–64, 65, 257
thunniform locomotion, 29
ticks, 135
tinamous, 235
Titanoboa, 149
titanosaurs
Alamosaurus, 127
Argentinosaurus, 119, 122, 123
discovery of, 119, 122
evolution of, in South America, 127
locomotion, 232
Tornieria, 111
torpor stage, 65
trace fossils, 59, 228
Trachodon mummy, 181–82
tracks
at Cal Orck’o mountain paleontological site, 233
of dinosaurs, 160, 161
in Dinosaur Valley State Park, 228
importance of, 228
in Kugitangtau Mountains (Turkmenistan), 228–29
at Lommiswil, 230–31, 231(illus), 232
mega-track complex that extends from southern Peru to northern
Argentina, 233
 at Münchehagen, 230
narrow-gauge and broad-gauge trackways, 232
of sauropods, 228
weight of animal and, 229–30
See also locomotion
Trelew International Airport, 123, 124
Triceratops, 178(illus), 182(illus)
anatomy, 178–79, 180–81
first discovery of complete skeleton of, 194
first discovery of skeleton of, 184
in Hell Creek and Lance formations, 203
nickname, 178
as one of last dinosaurs outside avian lineage, 177
as prey, 179–80, 196
sales of skeletons, 178
species of, 223–24
valid and invalid species names of, 179
Triceratops horridus, 223
Triceratops prorsus, 223
Tristan Otto (Tyrannosaurus rex), 195
Trix (Tyrannosaurus rex), 195
troodontids, 210
trophic levels, described, 40–41
Tsidiiyazhi abini, 260, 261
tsunamites, 256
Tübingen Museum of Natural History, 82–83
Tuebingosaurus, 82–83
tuffs, described, 42–43
Tyrannosauroidea, 214
See also Tyrannosaurus; Tyrannosaurus rex
Tyrannosaurus
anatomy, 210–11, 212
behavior of, 215–16
bite force of, 150
dentition of, 69, 149
earliest ancestor of, 162
 extinction of, 22
“niche assimilation,” 211, 212
number of species of, 221–23, 224–25
“secondary metamorphosis” of, 210–11
Tyrannosaurus imperator, 223
Tyrannosaurus regina, 223
Tyrannosaurus rex, 176(illus)
anatomy, 217–18
behavior of, 218
early discoveries of, 194
feathers and, 213, 216
juveniles, 208, 209(illus), 210, 212, 216
meaning of name, 194
naming of, 194–95, 223
number of, 199–201, 203–4
as one of last dinosaurs outside avian lineage, 177
popular images of, 193
prey of, 179–80, 196, 200, 201–2, 204–5, 208–9
range of, 198, 200
range of movement, 217–18
relatives of, 198
sales of skeletons, 197
sexual maturity, 208
size of, 200, 209(illus), 215
size of babies, 208
strength of, 217
time on Earth, 199

U
underwater flight, 54–55
University of Berkeley (California), 199
University of Bonn, 104
Upper Jurassic southern hemisphere dinosaur sites, 126
upright posture, advantages of, 67, 83–84
“Urvögel,” 72, 107
See also Archaeopteryx
V
Vectis Formation, 163
Vegavis, 245–46, 261
vision. See eyesight
vivipary, 47, 49–50
von Branca, Wilhelm, 112
von Meyer, Hermann, 76–77, 106
von Quenstedt, Friedrich August, 77
von Weinberg, Arthur, 185
von Zittel, Karl Alfred, 145

W
water column, described, 35
waterfowl, 236, 244, 260, 261
Wealden Group (Isle of Wight), 159
Wessex Formation (England), 91
When Life Nearly Died (Benton), 18
Wiemann, Jasmina, 242, 243
Winkelhorst, Herman, 32
Wintrich, Tanja, 36
Wyoming Dinosaur Center Foundation, 186

Y
Young, Bonda, 45
Young, Tom, 45
Yucatán Peninsula (Mexico), 256
Yutyrannus, 213–14, 215

Z
Zhuchengtyrannus, 198
zircons, described, 42–43

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ISBN-13: 9780369749369

The Lost World of the Dinosaurs

First published as Großartige Giganten in 2023 by dtv Verlagsgesellschaft

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