Mesozoic Birds: Above the Heads of Dinosaurs

533 Pages·2002·14.7 MB·English

by  Luis M. Chiappe

Mesozoic Birds

Mesozoic Birds
Above the Heads
of Dinosaurs
EDITED BY
LUIS M. CHIAPPE
AND
LAWRENCE M. WITMER
University of California Press
Berkeley Los Angeles London
Title page illustration: Archaeopteryx lithographicaby
Charles Knight (courtesy Los Angeles County Museum of
Natural History).
University ofCalifornia Press
Berkeley and Los Angeles,California
University ofCalifornia Press,Ltd.
London,England
© by the Regents ofthe University ofCalifornia
Library ofCongress Cataloging-in-Publication Data
Mesozoic birds :above the heads ofdinosaurs / edited
by Luis M.Chiappe and Lawrence M.Witmer.
p. cm.
Includes bibliographical references and index.
ISBN 0-520-20094-2 (alk.paper)
1. Birds,Fossil. 2. Paleontology—Mesozoic.
3. Birds—Evolution. I. Chiappe,Luis M. II. Witmer,
Lawrence M.
QE871 .M47 2002
568—dc21 2001044600
Printed in China
        
         
The paper used in this publication meets the
minimum requirements ofANSI/NISO Z.-
(R ) (Permanence ofPaper).∞
To Pat
and to Patty and Sam

Contents
Preface ix
Part I The Archosaurian Heritage of Birds
1 The Debate on Avian Ancestry: Phylogeny, Function, and Fossils
LAWRENCE M. WITMER 3
2 Cladistic Approaches to the Relationships of Birds to Other Theropod Dinosaurs
JAMES M. CLARK, MARK A. NORELL, AND PETER J. MAKOVICKY 31
Part II Taxa of Controversial Status
3 The Enigmatic Birdlike Dinosaur Avimimus portentosus: Comments and a Pictorial Atlas
PATRICIA VICKERS-RICH, LUIS M. CHIAPPE, AND SERGEI KURZANOV 65
4 The Cretaceous, Short-Armed Alvarezsauridae: Mononykusand Its Kin
LUIS M. CHIAPPE, MARK A. NORELL, AND JAMES M. CLARK 87
5 Alvarezsaurid Relationships Reconsidered
FERNANDO E. NOVAS AND DIEGO POL 121
Part III The Mesozoic Aviary: Anatomy and Systematics
6 Archaeopterygidae (Upper Jurassic of Germany)
ANDRZEJ ELZANOWSKI 129
7 The Discovery and Study of Mesozoic Birds in China
ZHOU ZHONGHE AND HOU LIANHAI 160
8 Sinornis santensis(Aves: Enantiornithes) from the Early Cretaceous
of Northeastern China
PAUL C. SERENO, RAO CHENGGANG, AND LI JIANJUN 184
9 The Birds from the Lower Cretaceous of Las Hoyas (Province of Cuenca, Spain)
JOSÉ L. SANZ, BERNARDINO P. PÉREZ-MORENO, LUIS M. CHIAPPE,
AND ANGELA D. BUSCALIONI 209
10 Noguerornis gonzalezi(Aves: Ornithothoraces) from the Early Cretaceous of Spain
LUIS M. CHIAPPE AND ANTONIO LACASA-RUIZ 230
11 Skeletal Morphology and Systematics of the Cretaceous Euenantiornithes
(Ornithothoraces: Enantiornithes)
LUIS M. CHIAPPE AND CYRIL A. WALKER 240
12 Vorona berivotrensis, a Primitive Bird from the Late Cretaceous of Madagascar
CATHERINE A. FORSTER, LUIS M. CHIAPPE, DAVID W. KRAUSE,
AND SCOTT D. SAMPSON 268
13 Osteology of the Flightless Patagopteryx deferrariisifrom the
Late Cretaceous of Patagonia (Argentina)
LUIS M. CHIAPPE 281
14 Enaliornis, an Early Cretaceous Hesperornithiform Bird from England,
with Comments on Other Hesperornithiformes
PETER M. GALTON AND LARRY D. MARTIN 317
15 The Mesozoic Radiation of Neornithes
SYLVIA HOPE 339
16 A Review of Avian Mesozoic Fossil Feathers
ALEXANDER W. A. KELLNER 389
17 The Track Record of Mesozoic Birds and Pterosaurs: An Ichnological
and Paleoecological Perspective
MARTIN G. LOCKLEY AND EMMA C. RAINFORTH 405
Part IV Functional Morphology and Evolution
18 Bone Microstructure of Early Birds
ANUSUYA CHINSAMY 421
19 Locomotor Evolution on the Line to Modern Birds
STEPHEN M. GATESY 432
20 Basal Bird Phylogeny: Problems and Solutions
LUIS M. CHIAPPE 448
Contributors 473
Index 477
viii CONTENTS
Preface
Less than two years after the publication ofCharles Dar- formation about the closest relatives ofthe living lineages of
win’s Origin ofSpecies,the first skeletal specimen ofArchae- birds (Neornithes).Ifthe abundant discoveries ofthe s
opteryx was discovered—a nearly perfect evolutionary made a significant contribution to our understanding ofthe
“missing link”—and since that time,the origin and early his- early evolution of birds, the s brought an embarrass-
tory ofbirds have been prime topics ofdebate among natu- ment of riches. In just that decade alone, the number of
ral historians and evolutionary biologists (De Beer, ; species ofMesozoic birds probably tripled those discovered
Desmond,;Elzanowski,Chapter in this volume).Not in all previous years (Chiappe,).These fossils were un-
much later,the paleontological exploration ofthe American earthed all around the world (Chiappe,;Feduccia,;
West resulted in the discovery ofmultiple specimens ofHes- Chatterjee,;Padian and Chiappe,),although pri-
perornis, Ichthyornis, and their kin (Marsh,), toothed marily in China,Mongolia,Argentina,Spain,Madagascar,
birds that,although much less primitive than Archaeopteryx, and the United States.
strongly influenced discussions about early avian evolution. A volume entitled Mesozoic Birds faces the sometimes
For many decades,this handful ofMesozoic taxa was all we sticky problem ofjust what constitutes a “bird.”In our nar-
had.It was the only fossil evidence available for understand- row time plane ofthe present day,birds are so markedly dis-
ing this fascinating chapter of vertebrate history.In fact,it tinct from other vertebrates that our perception ofwhat is
was not until the early s that significant new data started or is not a bird is patently obvious.But the Mesozoic era
to fill the extensive evolutionary gap separating the basal witnessed the dawning ofbirds,or,in scientific parlance,the
birds discovered in the previous century.The first evidence evolutionary transition to birds.As a result,the line between
ofa previously unknown radiation ofMesozoic birds,Enan- bird and not-bird is often a fuzzy one,and there are many
tiornithes,was announced by Cyril Walker in (Chiappe taxa whose avian status is highly controversial.This volume
and Walker,Chapter in this volume),and soon other taxa deals with several ofthese controversial taxa:for example,
(e.g.,the Mongolian Gobipteryx) were recognized as part of Protoavis (Chapter ), Caudipteryx (Chapters , , and ),
this radiation (Martin,).In addition,the discovery in Avimimus(Chapter ),Mononykusand its alvarezsaurid kin
the s of some fossil-rich Early Cretaceous lake beds in (Chapters and ).Decisions about where to draw the line
Spain (Sanz et al.,Chapter in this volume) and intensified could be made on the basis of particular features,such as
collecting in previously underexplored sites in Spain (Chi- feathers or a furcula (wishbone),but,as it turns out,virtu-
appe and Lacasa-Ruiz,Chapter in this volume) and China ally all the attributes that characterize modern birds were
(Zhou and Hou,Chapter in this volume;Sereno,Rao,and acquired sequentially in the Mesozoic.
Li,Chapter in this volume) led to many other discoveries Thus,the line between bird and not-bird is essentially ar-
not much younger than Archaeopteryx. The two-dimen- bitrary and must be defined.In this modern phylogenetic
sional preservation of this Early Cretaceous avian diversity era,taxonomic definitions are based on relationships,and
limited anatomical studies, but three-dimensional bird we have adopted for this volume the convention ofregard-
skeletons began to be found in other Mesozoic fossil sites ing Aves (and the colloquial terms “birds”and “avian”) as
(e.g.,Molnar,;Chiappe,).At the same time,other pertaining to the group comprising the most recent com-
Cretaceous discoveries (e.g., Kurochkin, ; Olson and mon ancestor of Archaeopteryx and neornithine (“mod-
Parris,;Hope,Chapter in this volume) afforded in- ern”) birds and all its descendants (Witmer,Chapter in this
ix