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Vertebrate Evolution
From Origins to Dinosaurs and Beyond
Donald R. Prothero
Illustrations by Nobumichi Tamura
First edition published 2022
by CRC Press
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and by CRC Press
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CRC Press is an imprint of Taylor & Francis Group, LLC
© 2022 Taylor & Francis Group, LLC
Reasonable efforts have been made to publish reliable data and information, but the author and
publisher cannot assume responsibility for the validity of all materials or the consequences of their
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Except as permitted under U.S. Copyright Law, no part of this book may be reprinted, reproduced,
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Trademark notice: Product or corporate names may be trademarks or registered trademarks and are
used only for identification and explanation without intent to infringe.
ISBN: 978-0-367-65176-3 (hbk)
ISBN: 978-0-367-47316-7 (pbk)
ISBN: 978-1-003-12820-5 (ebk)
DOI: 10.1201/9781003128205
Typeset in ITC Leawood
by Apex CoVantage, LLC
This book is dedicated to my students who have gone on to great careers in
paleontology or are still preparing for their future careers. These include
such paleontologists as John Foster, Jonathan Hoffman, Karen Whittlesey,
Jingmai O’Connor, Joshua Ludtke, Val Syverson, Katherine Long, Patrick
Gillespy, Thein Htun, Daniella Balassa, Sara Olson, Casey Cleaveland,
Kristin Watmore, and Katherine Marriott. They have encouraged and
supported me and inspired me throughout my teaching career, and I am so
proud of their accomplishments. They are the future of our profession.
CONTENTS
Preface
Acknowledgments
CHAPTER 1: Introduction: Finding, Dating, and Classifying Fossils
How Do You Find Fossils?
Dating Fossils
Naming Fossils
How Do We Classify Animals?
Further Reading
CHAPTER 2: The Origin of Vertebrates
What Is a Vertebrate?
Our Kinfolk in the Sea
Getting a Head: The Vertebrates
Conodonts
Further Reading
CHAPTER 3: Jawless Fish
Fish in Armor
Heterostracans
Thelodonts
Anaspids
Osteostracans
Galeaspida
Where Did They Come From? Where Did They Go?
Further Reading
CHAPTER 4: Primitive Gnathostomes
Jaws
Placoderms
Arthrodires
Antiarchs
Chondrichthyans: Sharks, Rays, Skates, and Chimaeras
Acanthodians
Further Reading
CHAPTER 5: Osteichthyes: The Bony Fish
Fish Bones
The Age of Teleosts
Further Reading
CHAPTER 6: The Transition to Land: The Tetrapods
Lobe-Finned Fish
Invasion of the Land: The Tetrapods
Further Reading
CHAPTER 7: Tetrapod Diversify
Amphibians and Their Relatives
Temnospondyls
Lepospondyls
Lissamphibians
Further Reading
CHAPTER 8: Primitive Reptiles
Land Eggs and the First Amniotes
Parareptiles
Eureptilia
Turtles
Further Reading
CHAPTER 9: Back to the Sea: Marine Reptiles
Ichthyosaurs and Plesiosaurs
Ichthyosaurs
Plesiosaurs
Placodonts
Further Reading
CHAPTER 10: The Scaly Ones: Lepidosauria—Lizards and Snakes
Lepidosauria
Rhynchocephalia
Squamates
Snakes
Mosasaurs
Further Reading
CHAPTER 11: Ruling Reptiles: Archosaurs
Archosauria
Archosauromorphs
Rhynchosauria
Allokotosauria
Proterosuchidae
Mystery Reptiles: Choristoderes
The Crocodile Branch: Pseudosuchia
Phytosaurs
Aetosaurs
Ornithosuchidae
Poposaurs
Rauisuchians
Further Reading
CHAPTER 12: Crocodylomorphs
Crocodylomorphs: The Crocodiles and Their Kin
“Bunny Crocs”
Notosuchia: The “Southern Crocodiles”
Thalattosuchia and Dyrosauridae: Back to the Ocean
Neosuchia
Further Reading
CHAPTER 13: Pterosaurs
Ornithodira/Avemetatarsalia
Flying Reptiles: The Pterosauria
Pterosaur Anatomy
Pterosaur Evolution
Further Reading
CHAPTER 14: The Origin of Dinosaurs
What Is a Dinosaur?
Further Reading
CHAPTER 15: Ornithischians I: Origins and the Thyreophora
The Ornithischians
Early Ornithischians
Thyreophorans: Stegosaurs and Ankylosaurs
Roofed Lizards: The Stegosaurs
Turtle-Shell Dinosaurs: The Ankylosaurs
Further Reading
CHAPTER 16: Ornithischians II: Hadrosaurs and Marginocephalians
Neornithischia
Ornithopoda: The “Bird-Footed” Dinosaurs
The Hadrosaurs
Pachycephalosaurs: The “Boneheads”
Ceratopsia: The Horned Dinosaurs
Further Reading
CHAPTER 17: Sauropods: Long-Necked Giants
The Largest Land Animals
The Origin of Sauropods
Jurassic Park of the Sauropods
Diplodocoids
Macronarians
Titanosaurs
Size Matters
Sauropod Physiology
Further Reading
CHAPTER 18: Theropods: Carnivorous Dinosaurs
Theropoda
Early Theropods
Tetanurae: Carnosauria
Carnosauria: Spinosauridae
Carnosauria: Megalosauridae
Carnosauria: Metriacanthosauridae
Carnosauria: Allosauridae and Carcharodontosauridae
Tetanurae: Coelurosauria
Coelurosauria: Tyrannosaurs
Coelurosauria: Compsognathidae
Coelurosauria: Ornithomimids
Coelurosauria: Maniraptora: Therizinosaurs
Coelurosauria: Maniraptora: Oviraptorosauria
Coelurosauria: Eumaniraptora: Dromaeosaurs
Further Reading
CHAPTER 19: Birds: The Flying Dinosaurs
Birds Are Dinosaurs
Mesozoic Bird Evolution
The Cenozoic Radiation of Aves
Neognath Birds
Neoaves
Terror from the Skies
Terror Birds
Further Reading
CHAPTER 20: Synapsids: The Origin of Mammals
The Origin of Mammals
To Be a Mammal
Early Synapsids: “Pelycosaurs”
Later Synapsids: “Therapsids”
The Third Wave: “Cynodonts”
Further Reading
CHAPTER 21: Primitive Mammals: Mesozoic Mammals, Monotremes, and
Marsupials
Mesozoic Mammals
Morganucodonts
Docodonts
Monotremes and Their Relatives
Multituberculates
Eutriconodonts
Therian Ancestors
The Marsupials or Metatheria
Further Reading
CHAPTER 22: The Placental Explosion: The Mammals Diversify
Placentals
Xenarthra: Sloths, Armadillos, and Anteaters
Afrotheria
Further Reading
CHAPTER 23: Laurasiatheria I: Carnivores, Bats, Insectivores, and Their Kin
The Laurasiatheres
Insectivores
Chiroptera (Bats)
Pholidota (Pangolins)
Carnivorous Mammals
Creodonts
Carnivorans
Further Reading
CHAPTER 24: Laurasiatheria II: The Ungulates
Horns, Hooves, and Flippers
Artiodactyls
Perissodactyls
Miscellaneous Mammals with Hooves
Pantodonts
Further Reading
CHAPTER 25: Euarchontoglires: Rodents, Rabbits, Primates—And Humans
The Euarchontoglires
Glires
Rodents
Lagomorpha
Euarchonta
Tree Shrews
Colugos
Primates
Strepsirhini
Haplorhini
Hominoidea (Apes and Humans)
Human Evolution
Further Reading
Index
PREFACE
I’ve already written several books about fossils and paleontology, including
a college textbook in paleontology for upper-level undergraduates (Bringing
Fossils to Life, 2nd ed., 2013, Columbia University Press) and a popular book
about fossil collecting (Fantastic Fossils, 2020, Columbia University Press),
but I’ve never found a book that gives a good general introduction to
vertebrate history for the general reader and fossil enthusiast with high-
quality color illustrations, so this book is intended to fill that need. Most of
the books on this topic for the general reader are picture books with minimal
information content. Given the rapid increase in knowledge about fossil
vertebrates, and the many changes in old notions about vertebrates and how
they lived, all such books are already grossly out of date. Thus, I wrote this
book to reach the general reader and especially the fossil enthusiasts and
collectors out there, who may have some knowledge about science, but not
necessarily a college-level background in geology or paleontology. There is a
real need to go beyond the pretty picture books, and present the latest
information about extinct vertebrates at a level that the general reader can
follow, but also enough information for college students in paleontology to
learn about the topic as well.
For this reason, the book is written at an intermediate level. I do not assume
any background in vertebrate anatomy, or the methods of systematics and
classification. Of course, the concepts are completely modern and in line
with the current thinking in cladistics, but I try to avoid jargon and
excessive technical terminology as much as possible. I have tried to use
familiar anatomical terms wherever possible, so it can be read and
understood not only by the amateur fossil enthusiast, but also by students
taking an undergraduate course in vertebrate evolution that does not require
training in anatomy or systematics. This is a very challenging task, because
most of the topics discussed in the book require a more advanced
understanding of systematics or anatomy, but I have done my best. I hope
the reader will find the book comprehensible and yet up to date and
accurate, incorporating all the latest thinking and discoveries of these
amazing animals.
ACKNOWLEDGMENTS
I thank my longtime friend Dr. Charles R. Crumly for encouraging me to
write this book and Ana Lucia Eberhart at CRC Press/Taylor & Francis for
editorial assistance. I thank Sathish Mohan of Apex CoVantage for
supervising the book design and production. I thank the following scientists
for their helpful reviews on various chapters: A. Henrici, D. Grossnickle, B.
MacFadden, K.L. Marriott, S. Modesto, S.J. Nesbitt, J. O’Connor, K. Padian, S.
Persons, T. Stidham, X. Wang, and several anonymous reviewers for the
comments and corrections when my presentation needed updating. I thank
K.L. Marriott for a lot of editorial help.
I thank the great paleontologists who taught and inspired me, from my early
career contacts with Drs. Dave Whistler and J. Reid Macdonald, to my
formal education in paleontology with Drs. Michael Woodburne, Michael
Murphy, Malcolm McKenna, Gene Gaffney, and Bobb Schaeffer. Without
their instruction and guidance, I would have never had the career in
paleontology that I sought since I first got hooked on dinosaurs at age 4.
From that age until today (over 63 years now), I never gave up, despite the
difficult challenges of finding a career in this crowded profession.
Finally, I thank my amazing family: my incredible wife, Dr. Teresa LeVelle,
and my sons, Erik, Zachary, and especially Gabriel, who also may become a
paleontologist someday. They put up with my long months at the computer
writing this book.
1
INTRODUCTION
FINDING, DATING, AND CLASSIFYING FOSSILS
DOI: 10.1201/9781003128205-1
Being a paleontologist is like being a coroner except that all the
witnesses are dead and all the evidence has been left out in the rain for
65 million years.
—Mike Brett-Surman, 1994
HOW DO YOU FIND FOSSILS?
Many kids grow up fascinated with dinosaurs and other prehistoric
creatures. Some even start digging holes in their backyards or driveways
looking for dinosaur bones. Most give up and become discouraged, because
fossil bones are extremely rare, and found only in a few places on earth.
If you wanted to find fossils, where would you look? Why are certain rocks
and certain places on earth good for finding fossils, while others have none
at all? First, nearly all fossils are primarily found in one kind of rock, known
as sedimentary rocks. These are rocks that are made from the loose grains of
sand, gravel, or mud, or other particles that weather out of the hard bedrock
and are deposited in rivers or flood-plains or in the bottom of the ocean.
