The Princeton Field Guide to Pterosaurs

185 Pages·2022·24.7 MB·English

by  Gregory S Paul

The
PRINCETON
FIELD GUIDE to

PTEROSAURS
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The
PRINCETON
FIELD GUIDE
to

PTEROSAURS
GREGORY S. PAUL
Princeton University Press
Princeton and Oxford
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Copyright © 2022 by Princeton University Press
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Library of Congress Cataloging-in-Publication Data
Names: Paul, Gregory S., author.
Title: The princeton field guide to pterosaurs / Gregory S. Paul.
Description: Princeton : Princeton University Press, [2022] | Series: Princeton
field guides | Includes bibliographical references and index.
Identifiers: LCCN 2021015050 (print) | LCCN 2021015051 (ebook) |
ISBN 9780691180175 (hardback) | ISBN 9780691232218 (ebook)
Subjects: LCSH: Pterosauria—Identification.
Classification: LCC QE862.P7 P38 2022 (print) | LCC QE862.P7 (ebook) |
DDC 567.918–dc23
LC record available at https://lccn.loc.gov/2021015050
LC ebook record available at https://lccn.loc.gov/2021015051
British Library Cataloging-in-Publication Data is available
Editorial: Robert Kirk and Abigail Johnson
Production Editorial: Kathleen Cioffi
Jacket Design: Layla Mac Rory
Production: Steven Sears
Publicity: Matthew Taylor and Caitlyn Robson
Copyeditor: Laurel Anderton
This book has been composed in Goudy Old Style (introduction), ITC Galliard
Pro (species section and headings), and Optima (labels)
Printed on acid-free paper. ∞
Typeset and designed by D & N Publishing, Wiltshire, UK
Printed in Italy
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CONTENTS
Preface 6
Acknowledgments 6
Introduction
History of Discovery and Research 9
What Is a Pterosaur? 10
Dating Pterosaurs 15
Evolution of Pterosaurs and Their World 16
Extinction 27
After the Age of Pterosaurs 28
Biology 29
General Anatomy 29
Walking, Running, Climbing, Swimming 36
Pterosaur Pneumatics 43
Skin, Feathers, and Color 44
Flight 46
Respiration and Circulation 68
Feeding Apparatus and Digestive Tracts 69
Senses 70
Vocalization 71
Disease, Pathologies, and Injuries 71
Behavior 72
Brains, Nerves, and Intelligence 72
Social Activities 73
Reproduction 73
Growth 76
Energetics 77
Gigantism 79
Mesozoic Oxygen 82
Pterosaur Safari 83
If Pterosaurs Survived 84
Pterosaur Conservation, Keeping, and Consumption 84
Where Pterosaurs Are Found 85
Pterosaur Count 86
Using the Group and Species Descriptions 91
Group and Species Accounts
Pterosaurs 97
Rhamphorhynchoids 98
Pterodactyloids 123
References 177
Index 181
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PREFACE
If I were, at about age twenty as a budding paleoresearcher and Remaining frustratingly unresolved are the origins of ptero-
artist, handed a copy of this book by a mysterious time traveler, saurs. Unlike with birds, whose descent from advanced dino-
I would have been startled as well as delighted. The pages would saurs has become well understood in many respects, but like with
have revealed a world of new pterosaurs and ideas that I had bats, the early evolution of which remains little documented in
only a hint of, if any such ideas existed at all. My head would the fossil record, we do not yet have a detailed understanding of
have spun at the revelation that Pteranodon was not, as had been which archosaur group pterosaurs evolved from, or how.
calculated, the largest flying creature possible, but that Quetzal- Producing this volume has been satisfying in that it has given
coatlus was even more colossal, with the wingspan of a small me reason to illustrate the skeletons of almost all pterosaur spe-
human-carrying glider, and a weight exceeding that of land- cies for which sufficiently complete material is available. These
bound ostriches manyfold. Visually pleasing would have been have been used to construct the most extensive library of side-
the array of pterosaurs bearing fantastical head crests, including view life studies of these archosaurs of the air to date in print.
Tupuxuara, Thalassodromeus, Shenzhoupterus, Tapejara, Sinopterus, An advantage of producing large sets of rigorous profile-skeletals
Tupandactylus, and most especially the superelongated, psyche- is that the restorations can reveal information that would not
delic prongs of Nyctosaurus, which had long been thought to be otherwise arise. One pertinent example, first appearing here,
dully crestless. Equally surreal would have been the hyperelon- is the discovery of the differing head/body proportions of Pter-
gated, superslender head of Moganopterus, which looks like it is anodon, one species of which was startlingly big beaked. The
out of an avant-garde Looney Tunes cartoon. Then there would overall result is a work that covers what is already well over two
have been the weirdly twisted beak of Cycnorhamphus. Plus the centuries of scientific investigation into the group of tetrapods
novel formations, at least to my eyes and ears, of Dolomia di that inhabited the skies for up to 170 million years. Enjoy the
Forni, Nugget Sandstone, Zorzino, Tiaojishan, Yixian, Jiufotang, travel back in time.
Tangshang, Toolebuc, Bissekty, Densus Ciula, Csebanya, Jagu,
Santana, Crato, and Javelina. The sheer number of new ptero-
saurs would tell that an explosion in pterosaur discoveries and Acknowledgments
research, well beyond anything that had previously occurred,
and often based on new high technologies, marked the end of
the twentieth century going into the twenty-first. Many thanks to those who have provided the assistance over
Confirmed would be the long-held speculation that the years that has made this book possible, including Peter Well-
pterosaurs—often called pterodactyls by the public (but those nhofer, Wann Langston, Paul MacCready, Alec Brooks, Asier
are just a portion of the group)—were not reptiles limited to Larramendi, Kenneth Carpenter, Michael Brett-Surman, Frank
gliding and weakly powered flight, but highly energetic fliers like Boothman, Brian Andres, Kevin Padian, David Peters, Michael
their distant archosaur relatives the birds, as well as mammalian Habib, Sandra Chapman, David Martill, Robert Telleria, and
bats. And, like bats, pterosaurs were covered with fuzz. How others. I would also like to thank all those who worked on
pterosaurs got around when not in the air has long been this book for Princeton University Press: Robert Kirk, Abigail
contentious—trackways have shown that they often walked on Johnson, Kathleen Cioffi, Layla Mac Rory, Steven Sears, and
all fours like bats, but the situation is otherwise complicated. Namrita and David Price-Goodfellow.
Pterosaurs are often seen as living in warm Mesozoic climes,
but some at least visited chilly polar regions and high altitudes.
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I N T R O D U C T I O N
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Anhanguera
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HISTORY OF DISCOVERY AND RESEARCH
Pterosaur remains have been found by humans for millennia area were discovered, first by Othniel Marsh and soon after by
and may have helped form the basis for belief in mythical beasts, his equally notorious competitor Edward Cope, the often largely
including winged dragons. In the prescientific West, the claim complete skulls and skeletons of the classic, largest known ptero-
in the Genesis creation story that the planet and all life were saur—and future star of feature films—Pteranodon. With wings
formed just two or three thousand years before the great Egyp- spanning 6.4 m (21 ft), it was a monster of the skies. At the turn
tian pyramids were built hindered the scientific study of fossils. of the next century, the much less enormous Nyctosaurus was
The discovery and subsequent detailed illustration of the virtu- found in the same beds, but unfortunately, those specimens did
ally complete and articulated skull and skeleton of the small not include its amazing head crest.
Pterodactylus found in fine-grained Lagerstätte sediments of Ba- After the loss of Seeley early in the 1900s, the new century saw
varia in the late 1700s began modern pterosaur paleozoology an extended slowdown in the paleozoology of pterosaurs. Good
decades before the discovery of teeth and a few bones led to the specimens continued to emerge from Europe, especially from the
recognition of land-bound dinosaurs in the 1820s. The strange Bavarian quarries, but nothing particularly novel. The same was
pterodactyl skeleton posed a major problem for early science true for the American plains, where Pteranodon specimens with
because it was still thought that major extinctions had not oc- broader, more erect crests than the better-known, more backward-
curred. The creature had a hyperelongated single finger, which bladed structures showed up. The onset of the age of aviation did
was radically different from the feathery airfoils of birds and also inspire a pre–Great War study on the aerodynamics of the big
quite distinctive from the multifingered wings of bats. Nor did pterosaurs, which were not all that much smaller than the first
the very long, spike-toothed jaws belong to either group. airplanes. And in the 1920s, Tilly Edinger did innovative work
In the early 1800s, various researchers believed that the pecu- on the avian-like structure of pterosaur brains. In the 1950s a
liar fossil, as well as more remains found in the Bavarian quarries fossil trackway was attributed to a pterosaur, but this assignment
that would later produce the feather-winged dinobird Archaeop- was disputed and little was done with it. Overall, remarkably
teryx, belonged to flippered swimmers, or to mammalian bats, little effort was expended on better understanding of pterosaur
perhaps of a marsupial nature. However, in the first decade of biology and flight, and a similar disturbing lack of vigorous sci-
the 1800s, the famed comparative anatomist Georges Cuvier ence afflicted the study of dinosaurs during the same period. Like
realized that pterosaurs were a distinctive group that used the dinosaurs, pterosaurs were seen as an extinct group, good for get-
single elongated finger to support a flight membrane. The new ting crowds through museum doors, but not of great biological
German remains further showed that there were two major sub- importance. And their origins remained vexingly obscure. The
groups of pterosaurs—the long-tailed rhamphorhynchoids epito- two superwars and grim economic troubles did not help matters.
mized by Rhamphorhynchus, and the short-tailed pterodactyloids Again in parallel to dinosaur science, pterosaur science started to
epitomized by Pterodactylus. English sediments began to produce revive in the 1960s and 1970s. In China, despite extreme political
pterosaurs, some found by the famed early field paleozoologist strife, new pterosaurs such as Dsungaripterus were described.
Mary Anning, and others described by Richard Owen, who de- Meanwhile, in Argentina the remarkable Pterodaustro, which
cided that the ancient fliers were low-energy reptiles. In the later sported long, slender, reverse flamingo-like filter-feeding jaws, was
1800s, Harvey Seeley became the leading pterosaur expert. He revealed. Of much greater paleobiological importance was the
concluded that because pterosaurs had fully developed wings publication of the central Asian Sordes. The specimen was a typical
like birds and bats, they should have had similarly high meta- small pterosaur, but what was remarkable was that it included an
bolic rates and were therefore not classic reptiles. This led to extensive covering of fur-like fibrous insulation, solid evidence
a bitter dispute between Owen and Seeley. At about the same for an elevated metabolic rate. That boosted the hypothesis that
time, it was realized that fossil thecodonts, as well as crocodil- pterosaurs were energetic at the same time that this same idea
ians, dinosaurs, birds, and pterosaurs, are all archosaurs. was being applied to dinosaurs. Peter Wellnhofer conducted an
The pterosaurs coming out of Europe were generally small extensive examination of the anatomy and biology of pterosaurs,
forms, often very well preserved, from the Jurassic period of the German and otherwise. During this preliminary pterosaur
middle Mesozoic, but some less complete English fossils showed renaissance, attention was again directed toward giant pterosaur
that Early Cretaceous pterodactyloids were of considerable size, flight, with the conclusion that pterosaurs were superlight aircraft
competitive with and even exceeding the size of the largest living that wafted around in the light airs of the warm and calm Mesozoic.
birds. What caused some head scratching was that these remains The stunning event that truly ignited the second golden age
were too fragmentary to reveal the identity of what seemed like of pterosaur paleozoology was the announcement in 1975 of an
bizarre rounded snouts and other oddities. Pterosaurian gigan- extreme pterosaur, Quetzalcoatlus, from just before the end of the
tism was confirmed when the focus of pterosaur paleozoology Mesozoic in Texas. Although the early wingspan estimates were
shifted to the newly open plains of North America. In the de- too high by as much as a factor of two, it was soon settled that
posits of the Late Cretaceous seaway that had then covered the these ultrafliers were borne by wings spanning 10 to 12 m (33
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