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The Rise of Reptiles
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The Rise of Reptiles
320 Million Years of Evolution

Hans-Dieter Sues

Johns Hopkins University Press Baltimore

© 2019 Johns Hopkins University Press Special discounts are available for bulk purchases of this book. For more
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Library of Congress Cataloging-in-Publication Data

Names: Sues, Hans-Dieter, 1956– author.

Title: The Rise of Reptiles: 320 Million Years of Evolution /
Hans-Dieter Sues.
Description: Baltimore : Johns Hopkins University Press, 2019. |
Includes bibliographical references and index.
Identifiers: LCCN 2018039554 | ISBN 9781421428673 (hardcover :
alk. paper) | ISBN 1421428679 (hardcover : alk. paper) |
ISBN 9781421428680 (electronic) | ISBN 1421428687 (electronic)
Subjects: LCSH: Reptiles—Evolution.
Classification: LCC QL645.3 .S84 2019 | DDC 597.9—dc23
LC record available at https://lccn.loc.gov/2018039554

A catalog record for this book is available from the British Library.
 Contents

Preface vii
Outline Classification ix

1 Introduction 1

2 Amniotes and Reptiles 15

3 Parareptilia: A Group of Their Own 22

4 Basal Eureptilia and Diapsida:

Early Evolution of Modern Reptiles 31

5 Testudinata: Turtles and Their Stem-Taxa 43

6 Sauropterygia, Ichthyosauromorpha, and Related Reptiles:

The Early Mesozoic Invasion of the Sea 72

7 Lepidosauromorpha: Rhynchocephalians, Squamates,

and Their Relatives 104

8 Archosauromorpha: The Ruling Reptiles and Their Relatives 159

9 Pseudosuchia: Crocodile-Line Archosaurs 177

10 Avemetatarsalia: Bird-Line Archosaurs Excluding Dinosaurs 209

11 Dinosauria I: Saurischia 229

12 Dinosauria II: Ornithischia 269

13 A Brief History of Reptiles 292

14 The Future of Reptiles 297

Glossary 303
References 309
Index 377
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Preface

Reptiles are one of the most diverse groups of present-day vertebrates, with more than
10,000 described species of crocodylians, lepidosaurs, and turtles. If one adds birds
(which are a clade of derived dinosaurs), this number more than doubles. Reptiles also
have a rich and varied fossil record spanning more than 300 million years. They in-
clude the largest land animals of all time, and they repeatedly and successfully invaded
the sea and took flight. An extraordinary wealth of new discoveries of extant and extinct
reptiles in recent decades has revolutionized our understanding of the evolutionary
history of these animals, but there exists no current overview. Carroll (1988) provided
the last major English-language review of this subject. Tatarinov (2006, 2009) attempted
a traditional synthesis of the evolutionary history of reptiles, which, in his usage, also
included synapsids excluding mammaliaforms. Finally, Benton (2014) published an ex-
cellent introduction to the evolution of reptiles in his textbook for beginning students.
Over the years several colleagues and students have asked me about a more current and
detailed exploration of this topic. The present work reviews the evolutionary history
based on our current knowledge. I hope that it will be of use to advanced students of
vertebrate zoology and paleontology.
Wherever possible, reptilian clades are phylogenetically defined and diagnosed in
terms of unambiguous shared derived features. Many taxonomic names have different
meanings for different researchers, and thus phylogenetic definitions of clades are crit-
ically important.
The illustrations include numerous color images of actual fossils to introduce readers
to the material basis for the study of extinct reptiles. In addition, numerous photographs
of present-day reptiles introduce paleontologists to the diversity of these animals. For
each major group, I have selected a phylogenetic framework or, where no compre-
hensive analysis was available, I have combined elements of compatible hypotheses. I
have tried to cite competing hypotheses for the (inter)relationships of par ticular reptil-
ian clades to alert readers to different points of view.
Anatomical terms follow the standard nomenclature used in comparative anatomy.
Stratigraphic terms follow the recommendations of the International Stratigraphic Com-
mission, and the absolute dates in Fig. 1.2 are based on the figures published by the
commission in 2018. Wherever possible, the periods, epochs, and ages for taxa are listed.
For the Permian Period, both the traditional and current tripartite divisions are listed.
Chapters 1 and 2 are introductory in nature. Chapters 3 through 12 review the di-
versity and evolution of the various reptilian clades. Chapter 13 presents a synopsis of
the evolutionary history of reptiles. Chapter 14 discusses the uncertain future of rep-
tiles in the context of the current global biodiversity crisis. A glossary provides the
viii Preface

definitions for numerous terms used in this volume. Robert Reisz, Olivier Rieppel, Rainer Schoch, and Xiao-
Finally, an extensive reference list invites readers to ex- chun Wu.
plore aspects of the evolutionary history of reptiles more I owe a great debt to friends and colleagues who gen-
deeply. It focuses on primary studies published in recent erously provided images for use in this book: Jérémy
decades because most of these publications include Anquetin, Sebastián Apesteguía, Sasha Averianov, the late
phylogenetic analyses of the taxa under consideration. Don Baird, Chris Bell, Mike Benton, Don Brinkman,
This does not reflect the common attitude today that the Chris Brochu, the late Alan Charig, Jim Clark, Adam
older literature is “outdated.” This body of work remains Clause, Ross Damiani, François Escuillié and the late Jean-
an indispensable source of primary data and citations. Claude Rage, David Evans, Mike Everhart, Nick Fraser,
Much to the dismay of many researchers, classifica- Greg Funston and Phil Currie, Heinz Furrer, Zulma Gas-
tion and nomenclature will always remain in flux as parini, Stephen Godfrey, Lance Grande, Harry Greene,
new studies on reptilian diversity and evolution contin- Romain Houssineau, Tom Jorstad, David Krause, Pete
uously provide new data and insights. The traditional Larson, Alexandra Laube, Stephan Lautenschlager, Jun Liu,
Linnaean system suggested that the relationships among the late Junchang Lü, Tyler Lyson, Jessie Maisano and
reptiles were far better resolved than is the case. It also Digimorph, Heinrich Mallison, Thomas Martens, Dave
created the misleading impression that categories such as Martill, Gerald Mayr, Ryosuke Motani, Sterling Nes-
“family” were somehow comparable actual biological enti- bitt, Bill Parker, Oliver Rauhut, Robert Reisz and Diane
ties. Thus, this system is not used in this book. I still refer Scott, Olivier Rieppel, the late Pamela Robinson, Rodolfo
to genera (as most genera of extinct reptiles comprise Salas-Gismondi, Torsten Scheyer, Rainer Schoch, Cesar
only a single species), but otherwise the higher-level tax- Schultz, Paul Sereno and Carol Abraczinkas, Adam Smith,
onomy in this book has dispensed with categories such Krister Smith, Juliana Sterli, Helmut Tischlinger, Mike
as family, order, and class. Like personal names, names Triebold, Andre Veldmeijer and Erno Endenburg, Laurie
of taxa are treated as singulars and without articles. Vitt, Mark Witton, Wolfgang Wüster, Pavel Zuber, and
Although this work is concerned primarily with the George Zug. Several museums and institutions kindly
fossil record, it makes frequent references to molecular- granted permission to reproduce photographs of speci-
based phylogenetic analyses. The latter have become the mens housed in their collections. Mike Ellison, Scott
principal tool for assessing the interrelationships among Hartman, Jeff Martz, and Paddy Ryan licensed use of
extant organisms. Morphology and molecules sometimes images created by them.
yield strikingly different phylogenetic hypotheses, but Special thanks are due to Zoe Kulik and Stuart Sumida
such conflicts lead to reconsideration of both kinds of who prompted me to embark on this project. At Johns
data sets. Hopkins University Press, Vincent Burke invited me to
write this book and, together with Tiffany Gasbarrini,
No single researcher can be familiar with all aspects of patiently saw this book through to publication. Susan
reptilian diversity and evolution I have tried to cover in Campbell provided superb copy-editing. I also thank
this book. Over the years I have learned a great deal from John Hoey and his team for their care during copy
friends through discussions and/or joint research. I par- production.
ticularly thank Sasha Averianov, Bob Carroll, Jim Clark, Last but not least, I thank Liz Sues for tolerating and
Nick Fraser, Sean Modesto, Ryosuke Motani, Sterling even encouraging my obsession with reptiles past and
Nesbitt, Paul Olsen, Kevin Padian, Adam Pritchard, present and for editorial comments on several chapters.
Outline Classification

This book uses a phylogeny-based classification of Reptilia. Relative ranking of clades is

indicated by indenting. Readers should consult Chapters 2 through 12 for further details.

REPTILIA
PARAREPTILIA
Mesosauridae
Unnamed clade
Millerettidae
Ankyramorpha
Lanthanosuchoidea
Unnamed clade
Bolosauria
Procolophonia
Pareiasauromorpha
Procolophonoidea

EUREPTILIA
Captorhinidae
Romeriida
Paleothyris
Diapsida
Araeoscelidia
Neodiapsida
Drepanosauromorpha
Weigeltisauridae
Testudinata
Testudines
Pleurodira
Cryptodira
“Mesozoic Marine Clade”
Ichthyosauromorpha
Hupehsuchia
Ichthyosauriformes
Ichthyopterygia
Ichthyosauria
Thalattosauriformes
x Outline Classification

Sauropterygia
Placodontiformes
Placodontia
Eosauropterygia
Eusauropterygia
Plesiosauria
Sauria*
(* denotes separate listing of the taxonomic categories within this clade below)

Phylogenetic classification of Sauria.

SAURIA
Choristodera
Lepidosauromorpha
Lepidosauria
Rhynchocephalia
Squamata
Iguania
Polyglyphanodontia
Unnamed clade
Mosasauria
Scleroglossa
Gekkota
Autarchoglossa
Scincomorpha
Lacertiformes
Amphisbaenia
Anguimorpha
Ophidia
Serpentes
Archosauromorpha
Crocopoda
Archosauriformes
Archosauria
Pseudosuchia
Suchia
Loricata
Crocodylomorpha
Crocodyliformes
Mesoeucrocodylia
Notosuchia
Neosuchia
Eusuchia
Crocodylia
 Outline Classification xi

Avemetatarsalia
Aphanosauria
Ornithodira
Pterosauria
Dinosauromorpha
Dinosauriformes
Dinosauria
Saurischia
Theropoda
Tetanurae
Avialae
Aves
Sauropodomorpha
Sauropoda
Ornithischia
Genasauria
Thyreophora
Stegosauria
Ankylosauria
Neornithischia
Marginocephalia
Ceratopsia
Pachycephalosauria
Ornithopoda
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The Rise of Reptiles
320 Million Years of Evolution
This page intentionally left blank
1 Introduction

A lifetime of fascination with, and many years of study of, reptiles past and present
has led me to writing this book. For several years I have reviewed all I have learned
in the course of my own research and surveyed the vast, widely scattered literature on
the subject. What follows is an attempt to present an overview of the evolutionary his-
tory of reptiles, from their modest beginnings to the diversity of species with which
we share this planet today.
There are more than 10,000 described present-day species of reptiles excluding birds.
This impressive, and still growing, number represents just three major lineages—turtles,
lepidosaurs (lizards, snakes, and the tuatara), and crocodylians (Fig. 1.1). Along with
birds (which, in phylogenetic terms, are a clade of feathered theropod dinosaurs), they
are the survivors of a once far more varied group of vertebrates.
For much of the past 300 million years or so, reptiles have been the most numerous
and diverse land-dwelling vertebrates on Earth. They repeatedly and successfully con-
quered the air and adapted to life in the sea. The Mesozoic Era, ranging from 251 to 66
million years ago (Fig. 1.2), represented the acme of reptilian abundance and diver-
sity and thus has long been known as the Age of Reptiles (Mantell 1831). One group of
reptiles, dinosaurs, included the largest land-dwelling animals of all time and another,
pterosaurs, included the largest flying animals that ever existed. Only after the extinc-
tion of dinosaurs other than birds at the end of the Cretaceous Period, some 66 million
years ago, did the major lineages of extant mammals (including our own) diversify.
Even today, however, reptiles including birds (with a combined total exceeding 20,000
described species) substantially outnumber mammals (about 6,500 species; Burgin
et al. 2018). Calling the Cenozoic Era the Age of Mammals merely reflects the fact that
we, a species of mammal, are writing the history of life.
Many years ago, Alfred Sherwood Romer, the leading vertebrate paleontologist of
the day, asserted that “the general pattern of reptilian evolution has become clear in
most regards” (Romer 1971:103). Fortunately for later generations of researchers, this
claim proved far off the mark. In recent decades, countless new reptilian fossils have
been discovered across the globe, both in previously unexplored regions and in well-
known locations. Combined with new approaches to the analysis of the interrelation-
ships of organisms and to the interpretation of fossil remains, these discoveries have
revolutionized our understanding of the reptilian Tree of Life.
Every year, driven by concerns about rapidly vanishing natural habitats around
the world, exploration efforts have led to the discovery of scores of new species of
present-day reptiles. Molecular methods—most recently, the sequencing of entire
genomes—provide critical new insights into the diversity and interrelationships of
2 The Rise of Reptiles

Figure 1.1. Representatives of the four major clades of present-day reptiles (excluding birds). A, Crocodylia: spectacled caiman
(Caiman crocodilus); B, Rhynchocephalia: tuatara (Sphenodon punctatus); C, Squamata: Boyd’s forest dragon (Lophosaurus boydii);
D, Testudines: Adanson’s turtle (Pelusios adansonii). A, courtesy of Laurie Vitt; B, courtesy of Paddy Ryan; C–D, courtesy of Division of
Amphibians and Reptiles, National Museum of Natural History.

extant species. This unprecedented wealth of new data exquisitely preserved fossil retains only a small amount
has clarified many issues but has also generated many of the total biological information the original life-
more questions. Thus, this book can provide only a “snap- form once did. Thus, fossils must be interpreted in the
shot” of our current understanding of the evolutionary context of what we know or can learn about present-
history of reptiles. day organisms.
Fossils are the remains or traces of ancient life. “Fossil”
Fossils derives from the Latin verb fodere, to dig. In the case of
Fossils are one of the principal sources of information for reptiles, fossils are typically bones and teeth, which are
the study of organic evolution. Although impressive in already fully mineralized in the living animal and thus
its own right, the diversity of present-day multicellular more likely to survive the death and dissolution of their
organisms represents only a tiny fraction of all living be- owner’s body. Only in exceptional circumstances have
ings that have ever existed on Earth—by some estimates soft tissues been preserved as impressions or become
as little as 1 percent (Nee and May 1999). Without fossils, rapidly mineralized soon after an animal’s death. Some-
our understanding of the diversity and history of life times trackways and other traces of life activity (such
would be extremely limited. However, even the most as eggs and burrows) can be confidently attributed to
 Introduction 3

actions with the surrounding sediment take place. Oc-

casionally, this leads to rapid precipitation of minerals
such as calcium carbonate around the remains, encasing
and protecting them from further destruction.
Fossils have established that there were once untold
numbers of animals and plants unlike anything alive
today. These extinct life-forms frequently show characters
or combinations of features not found in any present-day
species. Sometimes, fossils reveal that similar characters
evolved independently in different groups of organisms.
Fossils also help establish a time frame for evolution-
ary trees by providing minimum ages for the divergence
of present-day lineages, and they show us that many
groups of organisms were once much more diverse than
today in terms of number of body plans and species and
had different geographic distributions. Finally, fossils pro-
vide important data concerning the countless changes in
climate and the ever-changing distributions of land and
sea throughout Earth’s long history and the effects of
these changes on evolutionary events.
Reasonably complete skeletons of extinct reptiles are
not common. The spectacular skeletal mounts of dino-
saurs and other large extinct reptiles on display in natu-
ral history museums represent examples of exceptional
preservation or reconstructions based on (often multiple)
fragmentary specimens. Even in places where reptilian
Figure 1.2. The major divisions of the Phanerozoic Eon. fossils are abundant, the remains are commonly isolated
Radiometric ages as of 2018. Nomenclature and color scheme bones and teeth. Yet, to the trained eye, such remains re-
are based on the standards of the International Commission
veal a wealth of information about the reptiles to which
on Stratigraphy.
they originally belonged. Bones and teeth record changes
both subtle and gross during the life of an animal. So-
particular groups of extinct reptiles and provide impor- phisticated analytical methods for studying the micro-
tant additional insights into their biology. structure and chemical makeup of skeletal elements con-
The likelihood that an individual organism will be- tinue to provide new information concerning the growth,
come a fossil is vanishingly small. As soon as an animal diet, and health of their former owners.
has died, most of its bodily substance is quickly recycled. In a few extraordinary occurrences, known as conser-
Decay and dissolution of the soft tissues start immedi- vation fossil Lagerstätten, particular depositional condi-
ately, nourishing legions of microorganisms. Scavengers tions have led to the preservation of complete, articulated
both large and small will feed on the remains and often reptilian skeletons, frequently retaining even traces of soft
scatter what is left. Only if an organism is rapidly buried tissues. The fossils of the Paleogene (Eocene) strata of the
after death is there any chance that its hard parts will be Messel Pit in Germany provide a particularly spectacular
preserved. Even then, the chemical conditions in the sed- example, with mammals preserving hair and gut contents
iment have to be just right to keep those hard parts in- and birds preserving feathers (some even retaining traces
tact, or they too will eventually dissolve and vanish. As of their original color) and gut contents (K. T. Smith et al.
the soft tissues decompose, complex chemical inter- 2018). Such occurrences offer unmatched insights into the
4 The Rise of Reptiles

evolutionary history of many groups of animals and patience but can yield exquisitely preserved specimens.
plants and occasionally preserve life-forms for which Occasionally, bones have largely weathered away or are
there is little or no fossil evidence anywhere else. soft and crumbly in hard rock. In such cases, any remain-
The quest for fossils in the field requires patience and ing bony substance is carefully removed, leaving a sharp
luck. First, locations of potentially fossil-bearing sedimen- impression in the rock matrix. This impression is then
tary rocks are identified through study of the geological cast with a flexible molding compound (such as silicon
literature and maps. Then the real effort begins—looking rubber) to create a copy of the shape and surface texture
for and prospecting exposures of rocks. Some regions of the original fossil.
of the world have vast open terrain with little or no veg- Darwin (1859:310–311) noted despairingly, “I look at
etation and rock exposures everywhere. These are ideal the natural geological record, as a history of the world
hunting grounds for fossils. By contrast, searching for imperfectly kept, and written in a changing dialect. . . .
fossils in densely settled or vegetated areas like eastern Only here and there a short chapter has been preserved:
North America is much more challenging. There, out- and of each page, only here and there a few lines.” Al-
crops of fossil-bearing rocks are confined to quarries, though the fossil record has grown immeasurably since
road cuts, construction sites, riverbanks, and sea cliffs. Darwin penned these lines, parts of it remain tantaliz-
When visiting a potential fossil locality, collectors first ingly incomplete. Certain intervals of Earth history
scan the rock face for any bones or teeth already exposed apparently left little if any fossil record. However, pale-
by wind and rain. Occasionally such pieces will lead to ontologists have not yet fully explored many regions of
more complete skeletal remains still buried in the rock. the globe, and even historical localities continue to
At other sites, collectors must split rock to find fossils yield important new discoveries.
embedded in it. When smaller bones and teeth occur in
loose sediments, such as sands and clays, screening dry Molecules
or wet sediment through sets of sieves with a range of Biological molecules form the other major source of in-
mesh sizes is the most effective method used to recover formation for reconstructing the evolutionary history of
these remains. organisms. All living beings are made up of proteins. Pro-
Larger fossils are usually exposed to the degree nec- teins, in turn, are composed of chains of smaller mole-
essary to determine their extent and position in the cules known as amino acids. There are 20 common types
rock. They are then encased in protective jackets of of amino acids that can be assembled in any order to form
plaster-soaked bandages for safe transport back to a labo- a tremendous variety of proteins. The instructions for
ratory, where preparation—the actual excavation of the assembling proteins are encoded in a highly complex
bones and teeth—takes place. Standard tools for this molecule, deoxyribonucleic acid (DNA). DNA is a long
work include a kind of miniaturized jackhammer molecule with a “backbone” composed of alternating
(airscribe), chisels, dental picks, and sharpened rods of phosphate and sugar subunits. Attached to each sugar
steel or tungsten carbide mounted in pin vises. Detailed subunit is one of just four kinds of bases (nucleotides): ad-
preparation work is done with the aid of microscopes. enine, cytosine, guanine, and thymine. A set of three
Fossil bones are commonly fragile and riddled with bases (triplet) represents a single amino acid in a protein.
fractures, requiring consolidation as well as repair of A chain of such triplets makes up a gene. Over time, bases
cracks. Skeletal remains embedded in certain kinds of are changed, inserted, or lost, creating genetic mutations.
rock, especially limestone, can often be completely ex- Such mutations can be harmful by disabling a protein.
tracted through repeated immersion of the rock in greatly However, many, perhaps most, mutations appear to do
diluted acetic or formic acid. Limestone, composed pri- little if any harm, presumably because the genetic code
marily of calcium carbonate, readily dissolves in these includes much redundancy, and many regions of it do not
acids, exposing bones and teeth, which are made up of a code for proteins.
different calcium mineral, hydroxyapatite, and thus do Species diverge from common ancestors through
not dissolve as readily. This technique requires skill and changes in their DNA. By comparing the structure of the
 Introduction 5

DNA molecules between different species, biologists can codylians and turtles have the slowest rates of molecu-
track changes in the nucleotide sequences that have lar change reported among tetrapods, and snakes have
occurred since the divergence of those species from their some of the fastest (Hugall et al. 2007). Averaging rates
most recent common ancestor. Assuming that such of change across lineages in many molecular studies
changes took place at fairly constant rates over long spans pulls slowly evolving groups toward one another and
of time, researchers can estimate the amount of time it pushes rapidly evolving taxa toward the base of the
took for the differences to develop and compare the re- phylogenetic tree. Thus, much of the deep history of evo-
sults to the known fossil record. For example, humans lutionary lineages can probably not be discovered using
and chimpanzees share some 96 percent of their DNA. molecular methods.
Calculations for the time of the divergence of the human Ideally, the comprehensive study of the evolutionary
and great-ape lineages have estimated dates between six history of any group of organisms should draw on as
and seven million years, which fits well with the geologi- many sources of information as possible: molecules,
cal ages for the oldest known fossils of human-like pri- morphology, behavior, and ecology (total evidence ap-
mates from Africa. DNA sequencing has become the pri- proach; Eernisse and Kluge 1993).
mary tool for reconstructing the interrelationships of
present-day (and a few geologically young) organisms. Establishing the Age of Fossils
Much has been made of the fact that molecular and Establishing the absolute ages of fossils is essential for
morphological studies often yield strikingly different re- establishing time frames for the evolutionary diversifi-
sults. In such situations, potential issues affecting both cation of organisms. In most cases, however, it is impos-
sets of data must be carefully examined. Researchers fre- sible to date an individual fossil directly.
quently assume that molecular data are inherently supe- Only organic remains younger than about 60,000 years
rior to morphological information for reconstructing can be dated directly by using a radioactive isotope of
phylogenetic relationships. In most analyses of sequence carbon, carbon-14. Living animals and plants continu-
data, nucleotide sites (or amino-acid residues) are treated ously take up this relatively uncommon isotope from
as characters. In a large data set, such data can be hetero- the environment, but this uptake ceases when organisms
geneous. Another issue, known as long-branch attrac- die. Carbon-14 has a half-life of 5,730 ± 30 years, which
tion, emerges when evolutionary divergence between represents the period of time during which half of the
lineages under study occurred far back in time. If some original amount of carbon-14 has changed into nitrogen-14.
lineages have much longer temporal ranges than others, The amount of this isotope remaining can be measured
it is possible that these lineages evolved the same nucle- and provides a fairly accurate estimate of the age of a
otide at the same site because, after all, there are only four specimen.
possible nucleotides. Surprisingly, adding taxa to the By contrast, most of the fossils discussed in this book
analysis will compound this problem further rather than are much older—tens to hundreds of millions of years
resolve it (Felsenstein 1978). (Fig. 1.2). Although these fossils cannot be dated directly,
Molecular studies on present-day amniote tetrapods igneous rocks interlayered with the sedimentary depos-
sample only a few lineages in the Tree of Life—mammals, its containing the fossils can be dated using isotopes of
turtles, lepidosaurs, crocodylians, and birds. These various chemical elements with much longer half-lives
groups all diversified within the last 100 million years but than carbon-14. A widely used system involves two iso-
are connected to each other by lineages that ranged over topes of the noble gas argon, argon-40 and argon-39. Early
hundreds of millions of years (Z. Wang et al. 2013). attempts to determine the absolute ages of such rocks us-
Philippe et al. (2005) noted that molecular methods for ing isotopes were often fraught with large measurement
reconstructing phylogenies assume to at least some ex- errors, and reported ages varied considerably, but ad-
tent that rates of molecular evolution are broadly com- vances in methodology and analytical instrumentation
parable across lineages. In the case of reptiles, a number in recent years have made age determinations increas-
of studies have already challenged this assumption. Cro- ingly more consistent and precise.
6 The Rise of Reptiles

For most extinct species, determination of geological reconstruct the succession of reversals of Earth’s mag-
age relies on methods more indirect than radiometric netic field in geological time, establishing what is called
dating. Fossils occur in sedimentary rocks for which the the Geomagnetic Polarity Time Scale. Using the abso-
age can often be assessed by the presence of so-called in- lute ages of rocks for precise calibration, a regional mag-
dex fossils. Strata deposited at different locations often netostratigraphic succession can be correlated with this
yield fossil remains of the same species. If such taxa were global scale.
short-lived (in terms of geological time, existing for only
hundreds of thousands or a few million years), one can Linnaean Classification
determine that their host rocks were laid down within Classifying things is a basic human activity, and classi-
that particular time interval. The shorter the geological fying the natural world is no exception. Classification
range of a par ticu lar species, the more precisely the allows communication about life’s diversity and bestows
sedimentary deposits containing its remains can be a sense of stewardship. As common (vernacular) names
dated. In addition to being geologically short-lived, in- of animals and plants differ considerably even among
dex fossils should ideally be common and widespread. closely related languages, biologists long ago settled on a
Pollen and spores have proven particularly useful as universal system for classifying them to facilitate inter-
index fossils for dating continental sedimentary rocks. national scientific communication, employing the bi-
Plants produce them in enormous quantities. Wind or nominal system introduced by Carl Linnaeus in the eigh-
water often disperses these tiny propagules over great teenth century (Linnaeus 1758). In this system, organisms
distances; some even end up in marine deposits. Further- are grouped together in categories, known as taxa (singu-
more, many characteristic types of pollen and spores lar, taxon), within an explicitly hierarchical classifica-
came from plants that were apparently short-lived (in tion. The smallest commonly used taxonomic category,
terms of geological time) and widely distributed. This the species, bears a unique double name (binomen),
combination of attributes makes them particularly valu- much like our own personal names—most people have a
able index fossils. surname and at least one given name. In the case of spe-
Another increasingly impor tant tool for correlating cies, the equivalent of the surname is the genus name. A
rocks on a global level draws on changes in Earth’s mag- genus comprises two or more closely related species.
netic field over time. At the present time, the North The species name corresponds to the given name for a
Magnetic Pole is located close to our planet’s north rota- person. Each species is based on one particular speci-
tional pole—a condition known as normal polarity. For men, which is the holotype and provides the ultimate
reasons still not fully understood, Earth’s magnetic field standard of reference for that species. Linnaeus chris-
changes its polarity at irregular intervals, with the North tened our own species Homo sapiens, with Homo (Latin
Magnetic Pole moving close to the south rotational for “man”) being the genus name and sapiens (Latin for
pole (reversed polarity). Such reversals have occurred “wise”) the species name. Later researchers also referred
many times during the last 600 million years. They can to Homo several extinct species that proved closely re-
be traced in the rock record because certain minerals lated to anatomically modern humans. However, the
are easily magnetized. One such mineral, an iron oxide species name sapiens is restricted to the latter. Tradition-
known as magnetite, is common in various kinds of rock, ally, the names of genera and species were mostly derived
including basalts. As molten lava cools down to form ba- from Greek or Latin words because Latin was universally
salt, it passes through a threshold temperature known as used for scholarly communication in Linnaeus’s day,
the Curie point, the temperature at which magnetite and much as English is today. As individual genus names can
certain other minerals take up and lock in magnetiza- be employed only once, new names sourced from other
tion from Earth’s field. In sedimentary rocks, minute par- languages have become increasingly more common over
ticles of magnetic minerals become aligned with Earth’s time.
magnetic field when the sediments are deposited. Re- Linnaeus and his successors developed a formal hier-
searchers can measure this preserved magnetization and archy of nested sets of taxa to order the diversity of life:
 Introduction 7

genera were grouped into families, families into orders, relationships within that clade, and other features need
orders into classes, classes into phyla, and phyla into to be identified to sort out the internal structure of that
kingdoms. (Linnaeus himself originally used only class, clade. If the same or similar derived features are present
order, genus, species, and variety.) As our knowledge of in unrelated groups, they are considered the result of
the diversity of life grew, however, Linnaean classifica- independent, convergent evolution; such features are
tion continuously required introductions of additional known as homoplasies. Hennig’s approach is concerned
formal categories. In recent years, many researchers have with establishing the degree of relationship between taxa
begun to abandon these ranks altogether, retaining only rather than ancestry. Although species obviously have
species. ancestors, ancestry is scientifically untestable. For taxon
Linnaeus and his early followers classified organisms A to be the ancestor of taxon B, it must be demonstrated
based primarily on overall similarity. Many groups were that the former represents the inferred structure of the
recognized based on the absence of features: for exam- common ancestor of A and B in every detail. Not only is
ple, the vast majority of multicellular animals were this unlikely, but then there would be no shared derived
lumped together as invertebrates because they lack a ver- features to link the two taxa.
tebral column. To Linnaeus and other early students of In order to determine whether a particular feature is
the living world, the purpose of classification was to primitive (plesiomorphic) or derived (apomorphic) for a
bring order to nature’s vast diversity. set of taxa, it is compared with the condition in groups
that lie outside the set under consideration. For example,
Phylogeny and Classification when comparing features between different reptiles, a re-
Charles Darwin revolutionized scientific thinking about searcher would look at the corresponding traits in frogs
the history of life. He argued that species continuously and salamanders. Such groups are known as outgroups.
gave rise to other species over time. Following his lead, At first glance, an iguana and an alligator resemble
researchers moved from searching for features with each other much more closely than either resembles a
which to classify groups of organisms to exploring evo- chicken. The lizard and the alligator are usually grouped
lutionary links among groups. They sought to recon- together because they share many similarities, such as
struct detailed ancestor-descendant sequences and then scaly skin, typically sprawling limb posture, and a long
explained them in terms of adaptations to par ticular tail. However, these features are shared by most reptiles
modes of life. This led to the common but incorrect as- and represent primitive character states (which Hennig
sumption that paleontologists can somehow “read” pat- termed “plesiomorphies”) for that group. Closer exami-
terns of descent in the fossil record. nation shows that the alligator and the chicken share a
The work of Willi Hennig profoundly changed the number of derived features (synapomorphies), such as
way biologists examine how organisms are related to the presence of a muscular compartment of the stomach
each other. In a book first published in English in 1966, (gizzard), the complete division of the heart into four
Hennig argued that relationship is more meaningfully chambers, and the extensions of the air space in the re-
defined in terms of common ancestry rather than overall gion of the middle ear into the surrounding bones of the
similarity. His fundamental premise is that two taxa are cranium and mandible. They inherited these shared de-
more closely related to one another than either is to a rived character states from their most recent common
third only if they share a more recent common ancestor. ancestor. The iguana lacks these features. Thus, the alli-
If this is the case, the former two should share derived gator and the chicken are considered more closely related
features (which Hennig termed “synapomorphies”) that to each other than either is to the iguana.
they acquired from their most recent ancestor and that are When looking at the distribution of derived character
still absent in the third. If they do, the two taxa form a states in a group or groups of organisms, researchers of-
monophyletic or natural group, also known as a clade. ten find that more than one possibility can account for
Once synapomorphies for a particular clade have been the observed distribution of features. Each possibility in-
identified, they can no longer be used for determining volves different assumptions about how often the various
8 The Rise of Reptiles

features were acquired or lost. Following the principle to a clade that includes all the descendants of an event
of parsimony—the simplest explanation for the data is when a new group diverges from an ancestral stem. An
the most likely—scientists choose the arrangement that example is a stem-based definition of the dinosaurian
accounts for acquisition of the greatest number of fea- clade Saurischia as “all dinosaurs more closely related to
tures in the simplest way. The states of characters are Tyrannosaurus rex than to Triceratops horridus.” Finally, a
coded in numbers. The simplest example of such states taxon can be defined using a basal node. In the earlier
is the absence (scored as 0) or presence (scored as 1) of a example of Aves, a node-based definition could be “the
feature. For data sets with large numbers of taxa and most recent common ancestor of Archaeopteryx lithograph-
characters, computer algorithms assist in the search for ica and Passer domesticus (house sparrow) and all extant
the simplest arrangement. and extinct descendants of that ancestor.”
Hennig united taxa solely on the basis of derived fea- Wherever possible, clades should be defined by brack-
tures they share. Taxa are arranged into nested sets. In eting them with present-day species, which provide
illustrated diagrams, these sets are joined by lines that il- much more phylogenetically useful information than
lustrate the successive acquisition of features. A branch- even the best-documented extinct taxa. Crocodylia can
ing diagram in which all taxa occupy the end points of be defined as the most recent common ancestor of all
branches replaces the traditional family tree. Time is no present-day crocodiles, alligators, and gharials and all ex-
longer considered a critical factor for hypothesizing re- tant and extinct descendants of that ancestor. This node-
lationships. The key advantage of Hennig’s approach is based clade is firmly bracketed by extant species and
that researchers can continuously reassess old and add known as a “crown group.” Many Mesozoic crocodile-
new characters for groups of taxa and falsify or modify like reptiles were historically classified as crocodylians
hypotheses of relationships. based primarily on overall similarities. However, al-
Linnaeus and his followers classified organisms based though variously related to crown-group crocodylians,
primarily on the presence or absence of par ticular fea- these taxa lack some or even most of the features con-
tures. More recently, informed by Hennig’s approach, sidered diagnostic for the crown group. They are now
biologists began to define groups of organisms based on excluded from the crocodylian crown group and are
their ancestry. Under this approach, known as phyloge- referred to as “stem-crocodylians.” Some authors use
netic taxonomy and pioneered by Gauthier and de the concept of a total group (denoted by the prefix Pan-,
Queiroz (1990), an organism is assigned to a par ticular from Greek pas, all), which comprises a particular crown
group based on its place on the tree of life. Characters are group and all organisms more closely related to it than
used to reconstruct this tree, and shared derived features to any other crown group. For example, Pan-Aves com-
are employed to characterize, or diagnose, par ticular prises birds (Aves) and all taxa more closely related to
clades of organisms. In the previously discussed exam- them than to Pseudosuchia (Pan-Crocodylia), which com-
ple, the alligator and the chicken are assigned to a clade prises crocodylians and all taxa more closely related to
Archosauria (from Greek archon, ruler, and sauros, lizard), them than to birds (Gauthier and de Queiroz 2001).
which does not include lizards. Although increasingly widely used, the phylogenetic
Clades can be defined in various ways. The first def- approach to biological classification has been much crit-
inition is based on the presence of one or more apo- icized (e.g., Nixon and Carpenter 2000), primarily because
morphies. An example is a definition of birds (Aves) as of the considerable instability of classifications intro-
“all dinosaurs with wings and primary flight feathers.” duced by competing hypotheses of phylogenetic rela-
However, interpretations of features as primitive or de- tionships. However, biological classification has never
rived frequently change as new phylogenetic analyses been stable. It is important to remember that, as in any
draw on a broader range of taxa and new or revised char- other field of scientific inquiry, new discoveries contin-
acter assessments. Most researchers prefer other kinds uously test existing hypotheses. A biologically meaning-
of definition, which explicitly specify the basal member ful classification can never be fixed but should reflect the
of a particular clade. Stem-based definitions of taxa refer continually changing views of relationships.
 Introduction 9

present-day amniotes. This second phylogenetic analysis

Importance of Fossils for Phylogeny involved 67 “soft” features as well as 207 “hard” (skele-
Since the publication of Darwin’s magnum opus (Darwin tal) characters that could potentially be preserved in fos-
1859), fossils have been touted as the key evidence for sils. Many of these “hard” characters had previously been
evolutionary change. Given the spectacular advances in excluded from consideration because they were found
molecular biology in recent years, however, many biolo- only in par ticular present-day taxa and thus were not
gists have come to question the utility of fossils for re- helpful for elucidating the relationships between groups.
constructing the Tree of Life (e.g., Patterson 1981). One When Gauthier and his colleagues added fossil amniotes
example pertinent to this book underscores the enduring to their data matrix, many of these seemingly unique (au-
significance of fossils for this purpose. tapomorphic) traits emerged as more widely shared de-
Zoologists have long recognized that, among present- rived features. Again, present-day lungfishes (dipnoans)
day vertebrates, birds are most closely related to croco- and amphibians served as outgroups for identifying “soft”
dylians. This group is, in turn, most closely related to features as primitive or derived. By contrast, “hard” char-
lepidosaurs (lizards, snakes, and the tuatara), to the ex- acters could now be compared across a wide range of
clusion of mammals. Gardiner (1982) challenged this extinct tetrapods, some of which turned out to be more
orthodox view. Based on 37 derived features for extant closely related to amniotes than to either amphibians or
amniotes, he hypothesized that birds and mammals are dipnoans. In this second analysis, Gauthier and his col-
most closely related to each other and grouped them to- leagues recovered the traditional phylogeny of amniotes,
gether as Haemothermia (from Greek haima, blood, and with mammals (and stem-mammals) as the sister group
therme, heat; in reference to their warm-bloodedness). to reptiles including birds. Their study beautifully under-
Gardiner united Haemothermia and Crocodylia as Thec- scores the importance of fossils for reconstructing the
odontia and then grouped Thecodontia with turtles as evolutionary history of amniotes. Although imperfect,
Euamniota. fossils remain a key source of data for working out the
Gauthier et al. (1988a) tested Gardiner’s hypothesis. As evolutionary transformations of characters and thus for
a first step, they explored only the phylogenetic relation- testing relationships between taxa.
ships among extant amniotes. Gauthier and his col-
leagues critically assessed the characters compiled by The Skeleton of Reptiles
Gardiner (1982) and by a supporting study by Løvtrup Here follows a brief account of features of the reptilian
(1985). They modified some of these characters and their skeleton. For additional details, interested readers should
distribution among present-day tetrapods in light of new consult Romer’s (1956) classic Osteology of the Reptiles. Fur-
data and added new ones. Their revised data set had 109 thermore, the benchmark series Biology of the Reptilia
characters, most of which concern “soft” features that (1969–2010) contains a number of detailed reviews deal-
rarely if ever become fossilized. Present-day lungfishes ing with the skeleton of present-day reptiles.
(the closest relatives of tetrapods surviving today) and
amphibians served as outgroups for identifying particu- Skull
lar character states as primitive or derived. Gauthier and The skull consists of the cranium and mandible (Fig. 1.3).
his colleagues avoided using any information from fos- In the cranium, a set of bones surrounds the brain (brain-
sils in assessing character states for extant groups in the case) and is typically loosely attached to the elements of
phylogenetic analysis. The result of their analysis dif- the skull roof, palate, and sides. An opening for the nos-
fered from Gardiner’s study in recovering the tradition- tril (external narial fenestra or naris), the orbit (eye
ally recognized relationship between Crocodylia and socket), and (in most reptiles) one or two temporal open-
Aves, with Mammalia as the sister group to that clade. ings behind the orbit perforate either side of the cranium.
Gauthier et al. (1988a) then compiled a second data set, From the tip of the snout back, the side of the cranium
to which they added 24 tasa of extinct tetrapods as well comprises the following bones: the premaxilla and max-
as extinct representatives of the five major groups of illa, both of which usually bear teeth; the lacrimal and
10 The Rise of Reptiles

Figure 1.3. Structure of the skull of an early reptile, exemplified by the captorhinid eureptile Labidosaurus hamatus. Cranium in A,
dorsal; B, palatal; C, lateral; and D, occipital views. Lower jaw in E, lateral; and F, medial (lingual) views. Dermal bones, pale yellow
(rendered darker in deeper areas); bones of endochondral origin, orange; bones of the braincase, green; and stapes, pink. Abbrevia-
tions: an, angular; ar, articular; bo, basioccipital; bps, basisphenoid + parasphenoid; co, coronoid; d, dentary; ec, ectopterygoid; eo,
exoccipital; f, frontal; j, jugal; l, lacrimal; m, maxilla; n, nasal; op, opisthotic; p, parietal; pa, prearticular; pf, postfrontal; pl, palatine;
pm, premaxilla; po, postorbital; pp, postparietal; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sm,
septomaxilla; so, supraoccipital; sp, splenial; sq, squamosal; st, supratemporal; v, vomer. Courtesy of Sean Modesto and Robert Reisz.

prefrontal, which form the anterior margin of the orbit; orbit. The quadrate forms the jaw, or craniomandibular,
the postfrontal, postorbital, and jugal, which bound the joint with the articular bone of the lower jaw. Along with
orbit posteriorly and ventrally; and the quadrate, quadra- the epipterygoid, a rod-like bone that extends between
tojugal, and squamosal, which together make up the the palatal complex and the skull roof, the quadrate de-
“cheek” (temporal) region of the cranium behind the velops from the embryonic palatoquadrate cartilage.
 Introduction 11

The skull roof comprises (from front to back) the typ- either side of the cranium. Along the midline at the back
ically paired nasals, frontals, and parietals. The nasals of the palate, the parasphenoid covers, and is typically
usually separate the external narial openings, and the fused to, the ventral surface of the basisphenoid at the
frontals extend between the orbits along the midline of base of the braincase. Anteriorly, it forms a slender (cul-
the skull. In many reptiles, the parietals enclose between triform) process that extends forward in a space bounded
them a foramen, which is related to the pineal gland, a by the pterygoids (interpterygoid vacuity). In many rep-
structure of the brain that is sensitive to changes in am- tiles, the palatine, parasphenoid, pterygoid, and/or vomer
bient light and controls circadian rhythms. Basal reptiles bear small teeth or denticles.
retain additional pairs of bones—the postparietals, supra- The mandible comprises two rami (lower jaws or
temporals, and tabulars—along the posterior edge of the hemimandibles), which are typically connected anteri-
skull roof, but these elements are mostly or altogether orly by ligaments to form the mandibular symphysis. In
absent in more derived taxa. turtles and various other reptiles, the two rami com-
Most reptiles have openings in the roof and sidewall pletely fuse at the symphysis. The dentary is the prin-
of the cranium behind the orbits. These fenestrae, or tem- cipal tooth-bearing bone of each mandibular ramus.
poral openings, are related to the development of the Posteriorly, it contacts the surangular dorsally and the
adductor jaw musculature (Frazzetta 1968; Werneburg angular ventrally. The lingual (medial) surface of the den-
2012). When these jaw-closing muscles contract, they tary bears a groove for the Meckelian cartilage and is
generate considerable tensile stresses on the bones to covered by the splenial anteriorly and the prearticular
which they attach. Thus, muscles tend to attach to thick- more posteriorly. The coronoid bone forms a raised point
ened areas on bones such as ridges and crests. Develop- or process for the insertion of adductor jaw muscles and
ment of larger openings with rounded bony edges is also occasionally extends forward between the dentary and
thought to dissipate stresses because forces exerted by the splenial or prearticular. The posterior end of the man-
the jaw musculature can be distributed along the perim- dibular ramus is formed by the articular and bears a
eter of these openings. typically concave surface for contact with the quadrate.
On the back of the skull (occiput), the supraoccipital The hyoid apparatus supports the floor of the mouth
connects the posterior edge of the skull roof to the brain- and the tongue. Parts of this structure are sometimes cal-
case. Together with the paired opisthotics and exoccipi- cified or ossified and thus can be preserved in fossils.
tals, it typically bounds a large median opening in the The ear of most reptiles contains a single rod-like
occiput, the foramen magnum, through which the spi- bone, the stapes (columella auris), which transmits air-
nal cord passes from the brain posteriorly. The paroc- borne sound as vibrations from the eardrum (tympanic
cipital process of the opisthotic extends laterally toward membrane) to the inner ear. In some basal reptiles, how-
the bones of the temporal region. Together with the ever, the stapes is a robust element that probably served
prootic, and (when present) the laterosphenoid more an- primarily as a structural brace between the temporal
teriorly, the opisthotic forms the sidewall of the brain- region of the cranium and the braincase (Carroll 1980).
case. It also houses the semicircular canals of the inner The eyes of most extant and extinct reptiles have a
ear. Below the foramen magnum, the basioccipital and ring of thin bony plates (ossicles) embedded in the sclera
exoccipitals form the occipital condyle, which is part of around the pupil. This structure is thought to maintain
the joint between the head and neck (atlanto-occipital the shape of the eyeball or serve as the site of attachment
joint). The orbits are separated by a thin interorbital sep- for muscles for visual accommodation. As these scleral
tum, which occasionally is partially or fully ossified. ossicles are delicate, they are usually not preserved in
On the palate, the premaxillae make up the tip of the fossils.
snout, followed behind by the paired vomers and, further
posteriorly, the pterygoids, which extend posterolaterally Teeth
and contact the quadrates. The palatine and ectoptery- The free part of a tooth is the crown, and the part that
goid connect the pterygoid to the maxilla and jugal on is set in or attached to the jawbone is the root. Not
12 The Rise of Reptiles

referred to as presacral vertebrae. Each vertebra has a

body (centrum) and a neural arch on top of the centrum;
these two components surround the spinal cord and
are separate or fused together. The neural arch bears a
neural spine dorsally and is connected to the arches of
adjoining vertebrae by means of articular processes
known as zygapophyses. The vertebral centrum devel-
ops from tissue surrounding the notochord, a flexible
rod in the back of embryonic vertebrates, and the over-
Figure 1.4. Schematic transverse sections through jaws to lying neural tube. It replaces the notochord for the most
illustrate the traditionally recognized major types of tooth part. The centra of early reptiles often retain openings
implantation in amniotes. A, thecodont; B, pleurodont; and for the passage of a persistent notochord.
C, acrodont. Jaw, beige; tooth, blue; bony lining of the Unlike in many other tetrapods such as temnospon-
alveolus or attaching the tooth to the jaw, brown.
dyl stem-amphibians, the vertebral centra in reptiles are
large pleurocentra (Fig. 1.5). If present at all, the intercen-
surprisingly, in view of their varied dietary adaptations, tra are small, crescent-shaped bones wedged between
reptiles have a wide range of tooth shapes (Owen 1840– adjacent pleurocentra. The first two cervical vertebrae
1845; B. Peyer 1968; Edmund 1969; Berkovitz and Shellis are highly modified to facilitate motions between the
2017). Basal amniotes typically have teeth with simple head and neck. The first cervical or atlas (Fig. 1.6) is com-
conical crowns. In more derived reptiles, there are many posed of six distinct elements: the intercentrum of the
variations on this pattern. The upper and lower teeth atlas, which articulates below the occipital condyle; the
are usually spaced in such a manner that they do not pleurocentrum of the atlas; the paired neural arches of
come into contact with each other when the jaws are the atlas; and the paired proatlas. The atlanto-occipital
closed. Various reptilian groups, however, evolved tooth- joint permits rotation of the head on the neck. The sec-
to-tooth contact with tooth crowns modified for oral pro- ond cervical or axis has an intercentrum anteriorly, and
cessing of food. its pleurocentrum is fused to the large neural arch. The
Three basic types of tooth implantation are commonly cervical vertebrae typically bear ribs, which are shorter
recognized among reptiles (Fig. 1.4). Thecodont implan- and less massive than the ribs articulating with the dor-
tation refers to the teeth set in typically discrete sockets sal vertebrae. The dorsal ribs form the ribcage. In early
(alveoli) in the jawbone. Periodontal ligaments anchor reptiles, each rib head has two points of contact: the
and control limited movement of the teeth in the alve- capitulum contacts either the intercentrum or, more com-
oli. Acrodont implantation refers to the firm attachment monly, a facet (parapophysis) on the anterior edge of the
or fusion of the teeth to the apical ridges of the jawbones. pleurocentrum of the corresponding vertebra, whereas
Finally, pleurodont implantation reflects the reduction or the tuberculum articulates with a facet (diapophysis)
absence of the lingual bony wall of the jaw so that the on the transverse process of the neural arch. In some taxa,
tooth becomes attached only to the labial wall. There are the rib head forms only a single contact with the corre-
many variations on these three basic types. Proper char- sponding vertebra. Reptiles typically have at least two
acterization of the mode of tooth attachment often re- sacral vertebrae, each of which bears short, robust ribs
quires histological thin-sections or high-resolution CT connecting the vertebral column to the pelvic girdle on
scanning of jaw elements. either side. The tail usually comprises many vertebrae.
The centra of the caudal vertebrae bear Y-shaped ventral
Postcranial Skeleton bones known as hemal arches or chevrons. The two arms
The vertebral column is divided into (from the skull of the Y enclose blood vessels and nerves supplying the
back) neck (cervical), trunk (dorsal), sacral, and tail (cau- tail. Many groups of reptiles have rod-like bones known
dal) vertebrae. The cervicals and dorsals are collectively as gastralia that develop in the ventral body wall.
 Introduction 13

Figure 1.6. Structure of the atlas and axis of the basal diapsid
Petrolacosaurus kansensis in lateral view. Abbreviations: c1,2,
centrum of the axis and atlas, respectively; ic1,2, intercentrum
of the atlas and axis, respectively; na1,2, neural arch of the atlas
and axis, respectively; pa, proatlas. Modified from Reisz (1981).

glenoid forms a single screw-shaped surface, which re-

stricted the motion of the humerus at the shoulder joint
primarily to rotation about its long axis (Jenkins 1971).
In more derived groups, the scapula and coracoid each
bear glenoid facets, and the humerus has a much greater
range of mobility. The cleithra (if present) and clavicles
are thin, narrow bones extending along the anterior mar-
gin of the scapulocoracoid. The median interclavicle
Figure 1.5. Schematic structure of two dorsal vertebrae in connects the clavicles ventrally. The plate-like sternum
lateral view for A, a stem-amphibian, and B, a derived reptile. is positioned ventromedial to the ribcage. It is either
Neural arch, yellow; intercentrum, green; pleurocentrum, red;
single or paired but is not calcified or ossified in many
rib-head, white. A, modified from Mickoleit (2004).
reptiles.
Three pairs of major elements form the pelvic girdle:
Three pairs of major bones form most of each half the ilium dorsally, the pubis anteroventrally, and the
of the shoulder (pectoral) girdle: the scapula dorsally and ischium posteroventrally. The ilium is connected to
the coracoid (metacoracoid: Vickaryous and Hall 2006) the vertebral column by means of the sacral ribs. The
and procoracoid ventrally. They often fuse into a single pubis and ischium meet their opposites along a broad
scapulocoracoid. The scapula and coracoid form a com- symphysis ventromedially. Together with the ilium,
pound surface (glenoid) for articulation with the head they form a prominent articular surface (acetabulum) for
of the humerus. In early reptiles and their relatives, the contact with the femur.
14 The Rise of Reptiles

Each of the four limbs consists of a single proximal The manus primitively has five digits (fingers), each
bone (propodial) that articulates with two more distal comprising a metacarpal proximally and a series of
bones (epipodials), which in turn are connected with a smaller bones (phalanges) more distally. The phalangeal
series of smaller elements (mesopodials) that make up the formula for the manus in basal reptiles is 2-3-4-5-3, but it
wrist (carpus) or ankle (tarsus) and, more distally, the dig- is often considerably modified in more derived forms.
its (fingers or toes; autopodials). Within each digit, the The last phalanx or ungual of each digit typically bears a
bones embedded in the palm or sole are referred to as keratinous claw.
metapodials—metacarpals in the hand (manus) and meta- The hind limb comprises the femur in the thigh and
tarsals in the foot (pes). the tibia and fibula in the shank. The femur typically
The major bones of the forelimb are the humerus in bears distinct bony ridges or processes (trochanters) for
the upper arm and the radius and ulna in the forearm the attachment of thigh muscles. At the knee joint, it ar-
(antebrachium). The ulna extends lateral to the radius. It ticulates with the tibia, the principal bone of the shank
bears a large facet for articulation with the humerus and, (crus), and the fibula, which is situated lateral to the tibia.
above this joint surface, occasionally a bony olecranon The ankle (tarsus) comprises a proximal row composed
process for insertion of the triceps brachii muscle, which of the astragalus and calcaneum, an intermediate series of
extends the forearm. Several rows of small bones typi- centralia, and a row of distal tarsals. The astragalus origi-
cally form the wrist (carpus). The proximal row contacts nally formed by fusion of the tibiale, intermedium, and
the radius and ulna and comprises the radiale, inter- one or two centralia (O’Keefe et al. 2006). The pes in
medium, and ulnare. The distal row has five bones, one early reptiles has five digits (toes) with the same phalan-
contacting each digit. Between the proximal and distal geal formula as the manus, but again, more derived taxa
rows there is an additional set of bones, the centralia. show much variation in phalangeal counts.
2 Amniotes and Reptiles

The meaning of words tends to change over time, and “Reptilia” is no exception.
Linnaeus (1758) employed “Reptiles” (from Latin repere, to creep) for a subset of
limbed members of his “Amphibia” (which included various amphibians and reptiles
but also various fishes and the lamprey). Laurenti (1768) used “Reptilia” as a name for all
tetrapods other than birds and mammals. De Blainville (1816) first distinguished rep-
tiles from amphibians based on differences in the structure of their skins. Reptilian
skin is covered by scales, composed primarily of keratin, that protect the skin against
physical damage. Together with proteins and complex lipids in the epidermis, keratin
reduces the loss of water through the skin (Alibardi 2003). By contrast, present-day am-
phibians have moist skin that is rich in glands and much less keratinized than that of
reptiles. Gray (1825) also separated amphibians and reptiles, but most zoologists and
paleontologists adopted this distinction only later in the nineteenth century.
The most obvious difference between extant amphibians and other tetrapods is the
structure of their eggs. Amphibians typically lay eggs in fresh water. The eggs have a
clear, jelly-like coating. The developing embryo obtains oxygen for respiration and
most of its nourishment from the surrounding water and excretes its waste products
into the water. Most amphibians hatch as a larva, which continues to develop in
water and eventually undergoes metamorphosis to become an adult.
Extant reptiles, birds, and mammals have a different kind of egg than that of am-
phibians, the amniotic (or cleidoic) egg (Fig. 2.1). Haeckel (1866) first used the shared
possession of this feature to unite these three groups of tetrapods as Amniota (Fig. 2.2).
An amniotic egg develops membranes from “outside” the embryo (extraembryonic
membranes). One membrane encloses a large sac of yolk, which is connected to the
embryo’s digestive tract and provides nourishment. Folds of a membrane called the
amnion (from Greek amnos, lamb; the membrane was noticed in pregnant ewes) meet
and form a fluid-filled sac, the amniotic cavity, around the embryo. The sac-like allan-
tois grows from the embryonic hindgut and comes to surround the embryo. It serves
as a receptacle for metabolic waste products and aids in respiration. Another extraem-
bryonic membrane, the chorion, envelops almost the entire set of embryonic struc-
tures. This package, in turn, is enclosed in an outer shell, which can be either soft and
parchment-like or calcified. The shell is porous and connected through the chorion to
the allantois, which is well supplied with blood vessels and thus can take up oxygen
from and excrete carbon dioxide to the outside environment. The development of the
amniotic egg eliminated the need for a water-dwelling larval stage. In reptiles, a minia-
ture version of the adult hatches from the egg and makes its way in life. The acquisition
16 The Rise of Reptiles

and its replacement by a yolk sac. This change in the egg’s

coating would also have increased diffusion of oxygen
to the embryo. Simultaneously, the development of a po-
rous membrane around the egg would have strength-
ened it and further increased its permeability for respi-
ratory gases. These changes would have facilitated an
increase in egg size, which, in turn, would have required
provision of larger yolk resources for the embryo. In the
scenario proposed by Packard and Seymour, the yolk
sac was the first extraembryonic membrane to develop.
The allantois initially served only for storage of meta-
bolic waste products but subsequently increased in size
and assumed a key role in the exchange of respiratory
Figure 2.1. Simplified drawing of an amniotic egg in longitudi- gases.
nal section. Shell, dark gray; chorion, dark blue; amniotic To date no undisputed Paleozoic amniotic eggs have
cavity, light blue; embryo, green; yolk sac, yellow; allantois, been recovered. A possible example was reported from
pink. Modified from Bystrov (1957). the early Permian (Cisuralian) of Texas (Romer and Price
1939) but its identification as a shelled egg remains ques-
tionable (Hirsch 1979). The apparent lack of amniotic
eggs predating the Mesozoic Era is consistent with the
hypothesis that the eggs of early amniotes had parchment-
like shells, which would have rarely if ever become fos-
silized (Oftedal 2002). Parchment-like eggshell is still
found today in most lizards and in snakes. It much more
readily absorbs water than a mineralized shell (Oftedal
2002), and this attribute may have been important dur-
ing the initial evolution of the amniotic egg. As the
Euramerican region of Pangaea became increasingly drier
after the disappearance of the Pennsylvanian tropical for-
est communities (Sahney et al. 2010), water loss likely
became a critical issue for tetrapods. Mineralized egg-
shells would have limited water loss and increased the
eggs’ mechanical strength. They would have enabled
amniotes to venture from wet habitats into drier set-
tings and to diversify there, unlike other, more water-
Figure 2.2. Phylogenetic hypothesis of the interrelationships dependent tetrapods such as stem-amphibians.
of the major clades of present-day tetrapods, with coelacanths The amniotic egg acquires its shell within the female’s
(Actinistia) and lungfishes (Dipnoi) as outgroups. reproductive tract. As the egg must be fertilized before
the shell is formed, the male’s sperm must be introduced
into the female’s reproductive tract. By contrast, the
of the amniotic egg freed its producers from their de- males of most frogs and some salamanders simply shed
pendence on water for reproduction. their sperm on clutches of eggs deposited by the fe-
Packard and Seymour (1997) proposed a scenario for males in water. However, other present-day amphibi-
the evolution of the amniotic egg in which the first step ans internally fertilize their eggs. In most salamanders,
involved a reduction of the jelly-like coating of the egg for example, the female uses her cloaca to pick up a packet
 Amniotes and Reptiles 17

of sperm (spermatophore) deposited by the male. The

males of most extant (and presumably extinct) amniotes
have special intromittent organs for depositing sperm
inside the female’s reproductive tract. Turtles, croco-
dylians, some groups of birds (e.g., ratites), and mam-
mals have a median phallus. Lizards and snakes have
paired lateral hemipenes. The tuatara (Sphenodon puncta-
tus) lacks an intromittent organ, but Sanger et al. (2015)
demonstrated that its embryo has the paired tissue buds
from which the phallus develops in other amniotes.

Amniote Origins
It is not difficult to distinguish present-day amniotes
from amphibians. However, there has been a long-
standing debate whether various Pennsylvanian and
Permian tetrapods are early amniotes or even closely re-
lated to amniotes (Ruta et al. 2003; Klembara et al. 2014).
Traditionally, the origin of reptiles was linked to that
of the amniotic egg. However, fossils have since estab-
lished that the mammal and reptile-bird lineages had al-
ready diverged by the time the first amniotes appeared
in the fossil record during the Pennsylvanian. Thus, the
evolution of the amniotic egg predated this divergence,
and it is possible that even some stem-amniotes had
already evolved this type of egg.
The group generally considered most closely related
to amniotes is Diadectomorpha (Heaton 1980; Ruta et al.
2003; Reisz 2007). Diadectomorphs and amniotes are
grouped together as Cotylosauria (from Greek cotyle, cup,
and sauros, lizard; based on an early misinterpretation of
Figure 2.3. Skeleton of the stem-amniote Diadectes absitus in
the occiput) and share the absence of a distinct intertem- dorsal view. Courtesy of T. Martens.
poral bone in the skull roof (still present in earlier stem-
amniotes) and the shift of the postparietal and tabular
bones from the posterior portion of the skull roof onto Diadectes, from the Pennsylvanian and early Permian
the occiput. Cotylosauria also shares the fusion of the (Cisuralian: Gzhelian-Asselian) of the United States and
centra of the first two cervical vertebrae (atlas and axis), the early Permian (Cisuralian: Artinskian) of Germany,
the presence of broad, domed (“swollen”) neural arches attained a total length of up to 2.5 m (Olson 1947; Berman
on the dorsal vertebrae, and a sacrum comprising at least et al. 1998; Figs. 2.3, 2.4). Its skull has thick, porous bones,
two vertebrae. The phylogenetic analysis by Ruta et al. and it is often difficult to delineate individual cranial
(2003) recovered Amniota plus Diadectomorpha as most elements. The parietals are broad transversely. The quad-
closely related to a clade comprising Westlothiana, from rate is embayed posteriorly. Posteriorly, it contacts a thin
the Mississippian (Viséan) of Scotland, and Lepospon- plate of bone, which is part of the stapes (Berman et al.
dyli, a diverse but possibly not monophyletic group of 1998). The palatines form a partial secondary bony
mostly small-sized anamniote tetrapods ranging in time palate. The distinctive dentition of Diadectes consists of
from the Mississippian to the late Permian. forward-projecting (procumbent) incisor-like anterior
18 The Rise of Reptiles

Figure 2.4. A, skull and cervical vertebrae of the stem-amniote Diadectes absitus in dorsal view. Abbreviation: stp, ossified stapedial
plate. B, partial right dentary of Diadectes sideropelicus in occlusal view. Note incisor-like anterior and molar-like posterior teeth.
A, courtesy of Thomas Martens; B, from Case (1911).

teeth and molar-like posterior teeth with transversely pterygoid, which bears teeth in many forms and is re-
broad crowns (Fig. 2.4). The latter are frequently heavily lated to the differentiation of the pterygoideus jaw mus-
worn, and scratches on the wear surfaces indicate that cle. Another is the large and convex occipital condyle
the upper and lower teeth partially contacted each other that contacts a ring formed by the neural arch and inter-
and that the mandible moved fore-and-aft to break down centrum of the atlas; the centrum of the atlas contacts
food. The voluminous, barrel-shaped ribcage of Diadec- that of the axis (Sumida and Lombard 1991). This joint
tes indicates the presence of a capacious gut, which could between the head and neck permits a wide range of mo-
have accommodated endosymbionts to break down the tion. Finally, amniotes and various stem-amniotes
cellulose in the plant fodder. Diadectes and its close rela- share the possession of well-developed ungual phalan-
tives were among the earliest known tetrapods capable ges that are longer than the more proximal phalanges
of feeding on high-fiber plant matter (Hotton et al. 1997; and support keratinous claw sheaths in life.
Sues and Reisz 1998).
Another diadectomorph, Limnoscelis, from the Pennsyl- Amniotes
vanian (Gzhelian) of New Mexico and Colorado (Fig. 2.5), Amniota comprises the most recent common ancestor of
has (in plan view) a broadly triangular skull with large Synapsida (mammals and their close relatives) and Rep-
teeth at the tip of the snout (Berman et al. 2010). Its slig- tilia and all extant and extinct descendants of that an-
htly recurved tooth crowns have cutting edges close to the cestor (Reisz 1997, 2007; Fig. 2.6). Early amniotes differ
tip of the crown, suggesting that Limnoscelis was fauni- from other Paleozoic tetrapods in a number of skeletal
vorous. The girdles and limbs are robust. Limnoscelis at- features. On the skull roof, the frontal enters into the
tained a total length of about 2 m. dorsal margin of the orbit in basal amniotes, whereas
Some cranial features long considered diagnostic for it is excluded from that margin by the prefrontal and
amniotes (Carroll 1969a) are already present in various postfrontal even in the diadectomorph Limnoscelis. The
stem-amniotes (Ruta et al. 2003; Reisz 2007; Berman et al. well-ossified occipital condyle is hemispherical. Unlike
2010). One is the wing-like transverse flange of the in Diadectes, the posterior portion of the cranium in
 Amniotes and Reptiles 19

Figure 2.5. Reconstructed skeleton of the stem-amniote Limnoscelis paludis in lateral view. From Berman et al. (2010).

early amniotes lacks a distinct notch for support of the Scotia (Canada). Initially it was thought that these amni-
tympanic membrane. The ankle of amniotes includes a otes, along with other small animals, had become trapped
large astragalus, which formed through fusion of at and then entombed inside the hollow stumps. More re-
least three smaller bones (tibiale, intermedium, and cent work suggests that these animals probably lived in
one or two centralia). These constituent elements are the stumps or took refuge there from periodic wildfires
still evident on the astragali in some specimens of vari- (Falcon-Lang et al. 2010). The fossils recovered from the
ous Paleozoic reptiles (Kissel et al. 2002; O’Keefe et al. lycopsid stump casts include the oldest known reptile,
2006). Hylonomus, represented by disarticulated skeletons (Car-
As noted earlier, the skin of amniotes differs from that roll 1964). Geologically slightly younger Pennsylvanian
of extant amphibians. Not only is the epidermis itself (Moscovian) stump casts, also from Nova Scotia, have
more keratinized, but it is typically covered by scales, yielded articulated skeletal remains of the early reptile
feathers, or hair, all of which are also composed of kera- Paleothyris (Carroll 1969a) along with those of two taxa
tins. These features protect the skin against physical of basal synapsids. Carroll (1964, 1969a,b) interpreted
damage and limit water loss. At the same time, however, Hylonomus and Paleothyris as stem-reptiles broadly an-
this barrier greatly reduces the skin’s permeability for cestral to all later amniotes, but recent phylogenetic
oxygen and carbon dioxide. Thus, amniotes rely primar- analyses (Gauthier et al. 1988b; Müller and Reisz 2006)
ily on their lungs for breathing. Present-day sea snakes instead found these taxa closely related to diapsid rep-
and some turtles still are capable of limited cutaneous tiles (Chapter 4).
respiration, and water-dwelling turtles can even employ The absolute age boundaries for the Bashkirian are
their cloaca or pharynx to take up oxygen. Extant am- 323.2 and 315.2 Ma (Davydov et al. 2012). This is consis-
phibians, much like air-breathing fishes, push air from tent with the most recent estimate for the divergence of
their mouth and pharynx into the lungs. By contrast, am- crown Amniota with a minimum age of 318 Ma (Benton
niotes draw air into the lungs primarily by expanding et al. 2015). Based on these data, the book uses an esti-
the thoracic cavity, typically through movement of their mated divergence date of 320 Ma for the divergence of
ribs, which are linked by the intercostal muscles (Brain- Reptilia and Synapsida.
erd and Owerkowicz 2006). In most present-day am- There have been reports of geologically even older
niotes, each of the thoracic ribs is made up of multiple skeletal remains referable to amniotes. In the 1990s,
segments and articulates with the sternum. This enclo- Westlothiana was widely publicized as the earliest known
sure of the lungs by the ribs is necessary for breathing reptile. However, further preparation of the original find
by means of rib motion (costal aspiration). and the discovery of additional specimens have brought
The oldest undisputed fossils of amniotes to date have this interpretation into question. Westlothiana lacks some
been discovered in stumps of the tree-like lycopsid plant features such as the transverse flange of the pterygoid
Sigillaria from the Pennsylvanian (Bashkirian) of Nova that are present in amniotes and their closest relatives.
20 The Rise of Reptiles

However, it is certainly related to amniotes (Smithson have this condition. The other group, Diapsida (from
et al. 1994; Ruta et al. 2003; Clack 2012). Westlothiana Greek dyo, two, and hapsis, arch; in reference to the two
reached a snout-vent length of 11.5 cm. Although still bony arches bounding the temporal openings), typically
poorly known, Casineria, also from the Mississippian (Vi- has two fenestrae behind the orbit on either side of the
séan) of Scotland, is definitely a stem-amniote (Paton et al. cranium. A bony arch formed by the postorbital and
1999; Clack 2012). Its well-ossified postcranial skeleton squamosal bones separates the upper opening from the
has vertebrae with large pleurocentra, slender limbs, lower one, which itself is often bounded by a bony arch
and five-fingered manus with curved ungual phalanges. below. Crocodylians, lepidosaurs, and many extinct
Casineria attained an estimated total length of about 15 cm. groups of reptiles including stem-birds share this config-
uration, although it is considerably modified in many
Classifying Reptiles forms such as snakes and crown-group birds. Possibly for
The scientific exploration of extant and extinct reptiles ease of classification, Osborn also assigned turtles and
has made extraordinary strides in recent decades. Count- various other reptiles to Synapsida even though none of
less discoveries have uncovered a previously unimagined these forms have temporal openings. This led Williston
diversity of these animals spanning some 300 million (1917) to propose a third group, Anapsida (from Greek a,
years. This still-growing wealth of forms has continually without, and hapsis, arch; referring to the absence of bony
defied classification. arches), for reptiles that lack temporal openings. He
Osborn (1903) first distinguished two major groups interpreted the anapsid configuration as the primitive
among amniotes based on the structure of the skull. One condition for Reptilia. With changing definitions and
group, Synapsida (from Greek syn, together with, and varying constituents, the Osborn-Williston system of
hapsis, arch; in reference to the bony arch bounding the classification has remained in general use ever since (e.g.,
temporal opening), is characterized by the presence of a Romer 1956, 1966; Carroll 1988). However, many reptil-
single large opening (fenestra) behind the orbit on either ian taxa do not fit neatly into this system, repeatedly
side of the cranium. Mammals and their close relatives prompting proposals for additional groupings over the
(which, until quite recently, were classified as reptiles) years (e.g., Colbert 1945).

Figure 2.6. Phylogenetic hypothesis of the interrelationships of major clades of Reptilia. Diadectomorpha and Synapsida served
as outgroups. Testudinata, brown; Sauria, purple; Lepidosauromorpha, green; Archosauromorpha, red; “Mesozoic marine reptile
clade,” blue. Dots denote node-based clades and parentheses denote stem-based clades. Topology of the “Mesozoic marine reptile
clade” based on Scheyer et al. (2017).
 Amniotes and Reptiles 21

With the adoption of Hennig’s methodology for study- resolved. Thus, the definition of Reptilia becomes prob-
ing the relationships between organisms, the validity of lematic because, depending on whether turtles are or are
Reptilia was initially questioned. If mammals and birds, not part of Sauria, various taxa traditionally considered
historically treated as distinct classes in the Linnaean reptiles would be excluded from Reptilia. Modesto and
system, are separated from reptiles, “Reptilia” is reduced Anderson (2004) proposed a definition of Reptilia as the
to a grade of amniotes that lack the diagnostic features most inclusive clade containing Lacerta agilis (sand lizard)
of birds and mammals. Yet the name is historically clearly and Crocodylus niloticus (Nile crocodile) but not Homo
associated with, and has been in general use for, an as- sapiens (as a representative of Synapsida). This node-
semblage of present-day amniotes comprising turtles, based definition is adopted here.
crocodylians, and lizards and their relatives. As Reptilia The evolutionary lineages leading to mammals and to
is a more widely known name, it is given precedence over birds, respectively, diverged from each other early and can
Huxley’s (1869) Sauropsida for the clade comprising rep- already be distinguished by skeletal features among the
tiles and birds. Gauthier et al. (1988b) defined Reptilia as oldest known amniotes. The traditional division of Am-
the clade comprising the most recent common ancestor niota into three “classes”—reptiles, birds, and mammals—
of extant turtles and saurians (including birds) and all de- has given way to recognition of two major clades: Synap-
scendants of that ancestor. However, at that time, turtles sida (comprising mammals and their close relatives) and
were still considered most closely related to basal rep- Reptilia (comprising reptiles including birds) (Fig. 2.6).
tiles. The diapsid affinities of turtles are now well estab- The standard textbook statement that “mammals evolved
lished, but their position within Diapsida remains to be from reptiles” is phylogenetically not justified.
3 Parareptilia
A Group of Their Own

Olson (1947) proposed the name Parareptilia (from Greek para, beside, near, and Latin
repere, crawl) for a group comprising turtles, various late Paleozoic and early Meso-
zoic reptiles, and various taxa now considered stem-amniotes (e.g., Diadectes). He
noted that these forms share the presence of a well-developed otic notch, which is ab-
sent in other amniotes, which he named Eureptilia (from Greek eu, true, and Latin
repere, crawl). Olson’s hypothesis was at odds with then-prevailing ideas concerning
classification of reptiles and thus was largely ignored. Laurin and Reisz (1995) formally
resurrected the name Parareptilia for an assemblage comprising turtles and all amni-
otes more closely related to them than to diapsid reptiles. However, starting with
Rieppel and deBraga (1996) and supported by all molecular studies, most phylogenetic
analyses have hypothesized that turtles are related more closely to diapsids than to
parareptiles.
Tsuji and Müller (2009) defined Parareptilia (Fig. 3.1) as the most inclusive clade
containing Milleretta rubidgei and Procolophon trigoniceps but not Captorhinus aguti. Para-
reptiles first appeared in the fossil record in the Pennsylvanian (Modesto et al. 2015)
and became diverse during the Permian Period. Most are restricted to this period, but
one group, Procolophonoidea, survived the end-Permian extinction event and flour-
ished during the Triassic. Many parareptiles likely occupied ecological roles compara-
ble to those filled by eureptiles during later geological periods. Pareiasauria includes
the earliest known large-bodied reptilian herbivores on land. Reassessments of previ-
ously described taxa and numerous recent discoveries (e.g., Tsuji et al. 2010; Modesto
et al. 2015) hint at a much greater diversity of this clade yet to be discovered.
Parareptilia is characterized by a number of synapomorphies (Tsuji and Müller
2009): absence of caniniform teeth; postorbital region of the cranium short; posterior
margin of the skull roof forming a deep median embayment; jugal lacking a subtem-
poral process; and the absence of a supraglenoid foramen on the scapulocoracoid. The
crania of many parareptiles have a small lateral opening in or a distinct ventral emargi-
nation of the temporal region. Most have a posterior notch formed by the squamosal
and quadratojugal, which is conspicuous in taxa such as Macroleter and Procolophon
and probably supported a tympanic membrane in life. This notch, together with the
rather slender stapes, indicates that parareptiles evolved an impedance-matching ear
for receiving airborne sound independently from derived eureptiles and from mam-
mals (Müller and Tsuji 2007).
Several lineages of parareptiles evolved cranial and dental features that suggest
herbivory (Hotton et al. 1997; Reisz and Sues 2000; Reisz and Fröbisch 2014). This
change in diet independently occurred in a number of other amniote clades during
 Parareptilia 23

graphically well-defined distribution in an inland sea

on either side of the present-day South Atlantic Ocean
historically provided key evidence in support of the hy-
pothesis that Africa and South America were once con-
nected. Laurin and Reisz (1995) listed a number of shared
derived features for this clade including the absence of
caniniform teeth; the short postorbital region of the
skull; the absence of an ectopterygoid; retroarticular
process broad transversely and concave dorsally; and a
round proximal end of the femur. Mesosaurs attained a
total length of up to 1 meter. The greatly elongated, slen-
der jaws bear numerous, needle-like, pointed, and inward
curving marginal teeth. The more anterior teeth project
forward. The diet of mesosaurs consisted of crustaceans
Figure 3.1. Phylogenetic hypothesis of the interrelationships and occasionally small conspecifics (Silva et al. 2017). As
of Parareptilia. Dots denote node-based clades. Based on the delicate skull is usually badly crushed in known spec-
MacDougall and Reisz (2014).
imens, the presence (Piñeiro et al. 2012b) or absence
(Modesto 2006) of a lateral temporal opening remains
the Pennsylvanian and Permian. Feeding on plants re- uncertain.
quires various anatomical and physiological changes. The neck of mesosaurs is proportionately long. At
Vertebrates lack the requisite enzymes to break down least Mesosaurus and Stereosternum have accessory inter-
cellulose, which constitutes the bulk of the available vertebral articulations with zygosphenes and zygantra
plant material. Thus, herbivores have larger and more (Modesto 1999). Mesosaurs probably used the long later-
capacious guts housing vast numbers of endosymbiotic ally flattened tail and the hind limbs for swimming. The
bacteria and other microorganisms that can convert forelimbs are shorter than the hind limbs and may have
cellulose into volatile fatty acids, which can then be ab- assisted in steering. The structure of the elbow and ankle
sorbed by the hosts. The change in gut size is reflected joints indicates that mesosaurs likely had only limited ca-
by proportionately longer and/or broader rib cages. pacity for moving on land. The ribs in adults of Mesosau-
The breakdown of cellulose by endosymbionts is most rus are pachyostotic (with a thick periosteal cortex)
effective if plant fodder is already broken up prior to di- and “banana-shaped.” The thickening of the ribs and pre-
gestion. This is accomplished either by oral processing sacral vertebrae probably aided in controlling buoyancy.
or through the use of a muscular gizzard containing Pachyostosis is less extensive in other known mesosaurs
hard particles for grinding food (as in many extant (Houssaye 2009). Recent discoveries of embryos, includ-
birds). ing a tiny partial skeleton preserved inside the ribcage of
a medium-sized individual, show that mesosaurs were
Parareptilia: Mesosauridae viviparous (Piñeiro et al. 2012a).
Mesosauridae comprises the last common ancestor of
Mesosaurus, Brazilosaurus, and Stereosternum (Fig. 3.2) and Parareptilia: Millerettidae
all descendants of that ancestor (Laurin and Reisz 1995). Millerettidae comprises the most recent common an-
It represents the oldest known clade of amniotes adapted cestor of Milleretta and Millerosaurus and all descendants
to aquatic life. Modesto (2006) first placed Mesosauridae of that ancestor (modified from Laurin and Reisz 1995).
among parareptiles. Mesosaurs are known from the early It is known only from the middle to late Permian
Permian (Cisuralian: Artinskian) of Brazil, Uruguay, (Guadalupian-Lopingian: Capitanian-Wuchiapingian) of
Namibia, and South Africa (Oelofson and Araújo 1987; South Africa (Gow 1972, 1997). Diagnostic features in-
Modesto 1999, 2006; Piñeiro et al. 2012a,b). This geo- clude the lateral exposure of the quadrate, the sculpturing
24 The Rise of Reptiles

Figure 3.2. Skeleton of the mesosaurid Stereosternum tumidum. Courtesy of Robert Reisz and Diane Scott.

of the cranial bones consisting of low domed tuberosi- rate with nearly flat articular surfaces that are shorter
ties, and the cultriform process of the parasphenoid anteroposteriorly than wide transversely.
bearing denticles (Laurin and Reisz 1995). Milleretta at- Australothyris, from the middle Permian (Guadalu-
tained a total length of at least 20 cm and has dorsal ribs pian: Capitanian) of South Africa, has several diag-
with broad rather than rod-like shafts. The conical nostic cranial features such as a contact between the
crowns of the marginal teeth indicate a diet of arthro- postfrontal and supratemporal, a small interpterygoid
pods and other small animals. Gow (1972) reported the vacuity, and the presence of denticles on the ventral
presence of temporal openings in juveniles of Milleretta, surfaces of the basipterygoid processes (Modesto et al.
but these openings were obliterated in older individu- 2009a). It has a lateral temporal fenestra. Modesto
als. Millerosaurus has a ventrally open lower temporal et al. (2009a) considered the presence of this opening,
opening (Carroll 1988). It attained a skull length of about along with other derived character states such as the
5 cm. Eunotosaurus, from the middle Permian (Guadalu- deep posterior emargination of the quadrate, diagnostic
pian: Capitanian) of Malawi and South Africa, was con- for Procolophonomorpha.
sidered closely related to Millerettidae (Gow 1997), but Microleter, from the early Permian (Cisuralian: Sak-
several recent studies have reinterpreted it as an early marian) of Oklahoma, has a large number (32) of slen-
stem-turtle (see Chapter 5). der, slightly recurved maxillary teeth (Tsuji et al. 2010).
Its cranial bones bear ornamentation of tiny pits and
Parareptilia: Procolophonomorpha narrow radiating furrows. The cranium of Microleter
Procolophonomorpha (from Greek pro, before, kolophon, has a slit-like ventral emargination of the temporal
top, summit, and morphe, form) comprises the most recent region.
common ancestor of Australothyris and Ankyramorpha
and all descendants of that ancestor (Modesto et al. Parareptilia: Procolophonomorpha:
2009). Diagnostic derived features include the presence Ankyramorpha
of a lower temporal fenestra, a transversely broad Ankyramorpha (from Greek ankyra, anchor, and mor-
occipital condyle, and the distal condyle of the quad- phe, form; in reference to the shape of the interclavicle)
 Parareptilia 25

comprises the most recent common ancestor of Procolo-

phonia, Macroleter, Lanthanosuchidae, Acleistorhinus,
and all descendants of that ancestor (deBraga and Reisz
1996). This clade is diagnosed by several synapomor-
phies, notably the presence of a distinctly T-shaped
(“anchor-shaped”) interclavicle, which has long, slender
lateral processes bearing deep, forward-facing grooves
for the clavicles.

parareptilia: procolophonomorpha:
ankyramorpha: lanthanosuchoidea
Lanthanosuchus and its close relatives from the middle
Permian (Guadalupian: Capitanian) of Russia have broad,
dorsoventrally low crania with pronounced sculpturing
and lateral temporal fenestrae (Ivakhnenko 1987; Fig. 3.3).
The superficial similarity of this unusual cranial shape to
that in temnospondyl stem-amphibians initially led re-
searchers to exclude Lanthanosuchus and related forms Figure 3.3. Cranium of the lanthanosuchoid Lanthanosuchus
from amniotes, but more recent studies (e.g., deBraga and watsoni in dorsal view. Courtesy of Igor Novikov.
Reisz 1996) support parareptilian affinities for these pre-
sumably aquatic tetrapods. Lanthanosuchidae is most
closely related to Acleistorhinus, from the early Permian
(Cisuralian: Artinskian) of Oklahoma, which has a rather
box-like skull with a lateral temporal opening and with
sculpturing composed of shallow round pits (deBraga and
Reisz 1996). DeBraga and Reisz (1996) defined a clade Lan-
thanosuchoidea (from Greek lanthano, forget, and suchos,
crocodile) comprising the most recent common ancestor
of Lanthanosuchidae and Acleistorhinus and all de-
scendants of that ancestor. Diagnostic features listed by
these authors include the presence of a lateral lappet
Figure 3.4. Skull of the lanthanosuchoid Delorhynchus priscus in
of the frontal inserted between the prefrontal and post-
lateral view. Courtesy of Robert Reisz and Diane Scott.
frontal and contributing one third of the orbital margin,
the long basicranial articulation, and the absence of
basal tubera. Lanthanosuchoidea also includes Colobo- that at least this parareptile fed on arthropods (Modesto
mycter, Delorhynchus (Fig. 3.4), and Feeserpeton, all from et al. 2009b).
the early Permian (Cisuralian: Sakmarian) of Oklahoma
(Reisz et al. 2014). The teeth of these parareptiles have parareptilia: procolophonomorpha:
infolding of the dentine (plicidentine). Colobomycter has ankyramorpha: bolosauria
an enormous, tusk-like tooth in each premaxilla and a The closest known relative of Bolosauridae is Erpetonyx,
pair of much enlarged maxillary teeth; the remaining from the Pennsylvanian (Gzhelian) of Prince Edward
teeth are small. Feeserpeton has simple conical teeth, with Island, Canada (Modesto et al. 2015; Fig. 3.5). It is distin-
three enlarged ones in the maxilla and two in the den- guished by the presence of 29 presacral vertebrae and
tary. Some specimens of Delorhynchus preserved pieces lacks the specialized dentition of bolosaurids. Modesto
of chitin still adhering to their palatal teeth, confirming et al. (2015) placed Erpetonyx with Bolosauridae in a clade
26 The Rise of Reptiles

Figure 3.5. Skeleton of the bolosaurian Erpetonyx arsenaultorum.

Courtesy of Robert Reisz and Diane Scott.

Bolosauria (from Greek bolos, lump, and sauros, lizard).

Along with Mesosauridae, Erpetonyx demonstrates that
the initial diversification of parareptiles already occurred
during the Pennsylvanian.
Bolosauridae comprises the most recent common an-
Figure 3.6. A, reconstruction of the skull of the bolosaurid
cestor of Belebey, Bolosaurus, and Eudibamus and all de- Belebey vegrandis in lateral view; B, left lower jaw of Belebey
scendants of that ancestor. It is known from the early vegrandis in lateral view. Courtesy of Robert Reisz and Diane Scott.
Permian (Cisuralian: Asselian-Sakmarian) of Texas, Okla-
homa, France, and Germany and the middle Permian
(Guadalupian: Capitanian) of Russia and China (Reisz parareptilia: procolophonomorpha:
et al. 2007; Falconnet 2012). The dentition comprises pro- ankyramorpha: procolophonia
cumbent incisiform teeth in the premaxilla and anterior Following deBraga and Rieppel (1997), the name Pro-
part of the dentary and thickly enameled and bulbous colophonia is used for the grouping Pareiasauromorpha +
teeth in the maxilla and the posterior portion of the Procolophonoidea.
dentary that occluded with those in the opposing jaw
(Fig. 3.6). The maxillary teeth show heavy wear on the parareptilia: procolophonomorpha:
lingual surfaces of the crowns, whereas the dentary teeth ankyramorpha: pareiasauromorpha
are heavily worn on the labial sides. Well-developed stri- Tsuji (2013) found two synapomorphies for Pareiasauro-
ations on the wear facets extend more or less parallel to morpha: expanded epicondyle forming a broad, rectan-
the long axis of the jaw and indicate that the mandible gular, and anteriorly facing flange and astragalus and
was capable of fore-and-aft motion (Hotton et al. 1997). calcaneum fused to each other, with foramen for the per-
The lower jaw bears a tall coronoid process for attach- forating artery.
ment of the adductor jaw muscles (Reisz et al. 2007). Nycteroleteridae, including Macroleter (which is also
Bolosaurs were probably herbivores. The cranium has a known from Oklahoma) and Nycteroleter (Ivakhnenko
narrow lower temporal fenestra bounded ventrally by a 1987; Tsuji 2006), from the middle Permian (Guadalupian:
slender quadratojugal. Eudibamus, from the early Perm- Capitanian) of Russia, is characterized by a deep embay-
ian (Cisuralian: Artinskian) of Germany, attained a length ment in the posterolateral region of the cranium, which
of about 30 cm (Berman et al. 2000). It has hind limbs that is bounded by the quadrate, quadratojugal, and squamo-
are twice as long as the forelimbs and end in long, slen- sal and probably supported a tympanic membrane
der feet. Eudibamus is the oldest known reptile capable of (Müller and Tsuji 2007; Fig. 3.7). Macroleter also has a
bipedal locomotion, holding its hind limbs close to the small temporal opening between the jugal, quadratoju-
body and running on its toes. gal, and squamosal (Tsuji 2006). Nycteroleterids lack an
 Parareptilia 27

distinct shaft; and femur robust, with an only slightly

constricted shaft (Lee 1997a,b; Tsuji 2013). Some pare-
iasaurs attained total lengths of up to 3 m and weights of
at least 1,000 kg, making them the oldest known reptil-
ian megaherbivores. The cranium (Fig. 3.8) has a short
snout and a transversely broad postorbital region. Large
bony bosses are present on the “cheek” region of the cra-
nium, which extends below the tooth rows, and along
the ventral margin of the angular. The marginal teeth of
pareiasaurs have labiolingually flattened, leaf-shaped
crowns with coarsely serrated cutting edges (Lee 1997a)
and superficially resemble those of extant plant-eating
iguanid lizards. They are closely spaced and form contin-
uous shearing edges. The vertebral column of pare-
iasaurs comprises only 17 to 20 presacral vertebrae. The
massive, barrel-shaped rib cage indicates the presence of
a capacious gut for lengthy retention and fermentative di-
gestion of plant fodder (Reisz and Sues 2000). The pelvis
has a large ilium and a smaller ischium and pubis, which
are aligned behind the ilium in some taxa. The robust
Figure 3.7. A, cranium of the nycteroleterid Macroleter poezicus
in lateral view; B, skull and partial postcranial skeleton of
limbs of pareiasaurs were held more vertical than in
Macroleter poezicus in dorsal view. A, from Müller and Tsuji other parareptiles to support the massive body. M. L.
(2007)—CC BY 2.5; B, courtesy of Robert Reisz and Diane Scott. Turner et al. (2015) reconstructed Bunostegos, from the
Permian of Niger, with a more upright forelimb posture
than in other pareiasaurs that have more sprawling fore-
intertemporal (Lee 1995). They have skull lengths rang- limbs. The broad manus and pes have reduced numbers
ing from 5 to 10 cm. Tsuji and Müller (2009) and Tsuji
(2013) considered nycteroleterids most closely related
to pareiasaurs. Tsuji et al. (2012) united Nycteroleteridae
with Pareiasauridae in a clade Pareiasauromorpha.
Pareiasauridae (from Greek pareion, cheek, and sauros,
lizard) comprises the most recent common ancestor of
Bradysaurus and Anthodon and all descendants of that
ancestor. This clade ranged in time from the middle
to late Permian (Guadalupian-Lopingian: Capitanian-
Changhsingian) and is known from various regions of
Africa and from Brazil, China, Germany, Russia, and
Scotland (Ivakhnenko 1987; Lee 1997a,b; Jalil and Janvier
2005; Tsuji 2013). Laurin and Reisz (1995) provide a list
of diagnostic features including the exclusion of the fron-
tal from the orbital margin; sculpturing of cranial bones
comprising large knobs, pits, and ridges; presence of a
distinct ventral boss near the anterior end of the angu-
larr; three to five pairs of sacral ribs; scapula with a prom- Figure 3.8. Skull of the pareiasaur Deltavjatkia rossica in lateral
inent acromion; humerus short, robust, and lacking a view. Courtesy of Robert Reisz and Diane Scott.
28 The Rise of Reptiles

of phalanges. Many pareiasaurs bear extensive dermal ar-

mor (Lee 1997a,b; Tsuji 2013). In basal taxa such as Bradys- parareptilia: procolophonomorpha:
aurus, from the middle Permian (Guadalupian: Capitanian) ankyramorpha: procolophonia:
of South Africa, this armor is composed of thick, rounded, procolophonoidea
and coarsely sculptured bony plates (osteoderms) embed- This clade is supported by various derived features in-
ded in the skin along the dorsal midline. In more derived cluding the presence of a narial shelf, the posterior ex-
pareiasaurs, such as Anthodon, from the late Permian tension of the jugal that contributes to the temporal
(Lopingian: Wuchiapingian) of South Africa, and Scuto- region, a ventrally open lower temporal opening open
saurus, from the late Permian (Lopingian: Changhsin- ventrally, and three or more bones forming the retro-
gian) of Russia, the osteoderms are arranged in closely articular process (MacDougall and Reisz 2014).
spaced transverse rows across the back. Even the limbs Nyctiphruretidae is characterized by the presence of
bear small conical armor elements. paired surangular foramina (MacDougall and Reisz 2014),
and comprises Nyctiphruretus, from the middle Permian
(Guadalupian: Capitanian) of Russia, and Abyssomedon,
from the early Permian (Cisuralian: Sakmarian) of Okla-
homa. The cranium of Nyctiphruretus has a long external

Figure 3.9. Two associated skeletons of the owenettid Figure 3.10. Crania of procolophonids. A, cranium of the
“Owenetta” kitchingorum in dorsal view. The associated Early Triassic Procolophon trigoniceps in lateral view; B,
worm-like structures represent fossils of millipedes, which cranium of the Middle Triassic Teratophon spinigenis in dorsal
presumably scavenged the reptilian remains. Courtesy of Robert view. A, courtesy of Robert Reisz and Diane Scott; B, courtesy of
Reisz and Diane Scott. Ross Damiani.
 Parareptilia 29

narial opening bounded by a large depression (Säilä 2010a). ated. Furthermore, a few procolophonoids have a lower
Nyctiphruretus attained a total length of about 40 cm. temporal opening (e.g., Candelaria; Cisneros et al. 2004).
Procolophonoidea (from Greek pro, before, and kolo- Owenettidae ranged in time from the late Permian
phon, top, summit) comprises the most recent common (Lopingian: Wuchiapingian) to the Middle Triassic (La-
ancestor of Owenettidae and Procolophonidae and all de- dinian) and is known from Brazil, Madagascar, South
scendants of that ancestor (Lee 1997a). It ranged in time Africa, and Tanzania. Owenetta, from the late Permian
from the late Permian (Lopingian) to the end of the Tri- (Lopingian: Wuchiapingian) of South Africa, and its
assic and attained a worldwide distribution (Ivakhnenko relatives have conical, slightly recurved marginal teeth
1979; Cisneros 2008). Diagnostic derived features for this (Reisz and Laurin 1991; Reisz and Scott 2002; Fig. 3.9).
clade include the presence of a large medial process of the Diagnostic apomorphies include the presence of a deep
prefrontal; the elongation of the frontal; a distinctly temporal emargination between the jugal and quadrato-
domed and large supratemporal; the absence of the post- jugal and the separation of the postorbital and parietal by
parietal; and the absence of teeth on the transverse a large postfrontal (Reisz and Scott 2002).
flange of the pterygoid (Reisz and Scott 2002). The skull The more diverse Procolophonidae (Figs. 3.10, 3.11)
of procolophonoids is proportionately short anteropos- spanned the entire Triassic Period. The largest known
teriorly and broad transversely. The posterior margins of procolophonid, Sclerosaurus, from the Early to early
the orbits are deeply embayed and probably accommo- Middle Triassic (Olenekian-Anisian) of Germany and
dated part of the adductor jaw musculature (Figs. 3.8, 3.9). Switzerland, attained a total length of 50 cm (Sues and
The temporal region of the cranium is ventrally emargin- Reisz 2008), but most taxa are smaller. Cisneros (2008)

Figure 3.11. Skeleton of the

procolophonid Tichvinskia
vjatkensis in dorsal view,
probably originally preserved
in a burrow. Courtesy of Igor
Novikov.
30 The Rise of Reptiles

listed a number of synapomophies for Procolophonidae the Late Triassic of Scotland, the lower jaw has a tall,
including the absence of a premaxillary process of the sometimes recurved coronoid process, and the jaw joint
maxilla; maxilla with a lateral depression behind the ex- is placed below the level of the lower tooth row (Sues
ternal naris; three or four conical or incisiform premax- et al. 2000; Säilä 2010b). These features, along with the
illary teeth; and maxillary and more posterior dentary structure of the teeth, indicate herbivory for at least the
teeth with labiolingually broad crowns. When unworn, more derived procolophonids (Reisz and Sues 2000).
each molariform tooth in the maxilla and dentary typi- The quadratojugal in many procolophonids is either ex-
cally has a labial and a lingual cusp connected by a sharp tended posterolaterally or bears two or more prominent
crest. Opposing tooth crowns fit between each other bony spines; these features probably bore keratinous
like cogs when the jaws closed and could have crushed covering in life and may have served as protection
and sheared food trapped between them. In derived against predators or for some other purpose such as
procolophonids such as Hypsognathus, from the Late digging. Articulated skeletons of owenettids and pro-
Triassic (Norian-Rhaetian) of Connecticut, New Jersey, colophonids have frequently been found preserved in
Nova Scotia, and Pennsylvania, and Leptopleuron, from burrows (Ivakhnenko 1979; Figs. 3.9, 3.11).
4 Basal Eureptilia and Diapsida
Early Evolution of Modern Reptiles

Traditional scenarios of tetrapod evolution depicted reptiles as evolving from “amphibi-

ans” (i.e., anamniote tetrapods) and subsequently giving rise to birds and mammals, re-
spectively (Romer 1966; Carroll 1969a,b). The ancestral reptilian stock was thought to
comprise small-bodied, superficially lizard-like forms like the Pennsylvanian Hylonomus,
which have skulls without temporal openings. Reassessments of the interrelationships
of amniotes using Hennig’s methodology, however, found no support for this view. In-
stead, they established the existence of two major lineages of amniotes, one leading to
eureptiles (including birds) and the other leading to mammals, both of which first appear
in the fossil record during the Pennsylvanian (Gauthier et al. 1988a,b; Reisz 1997).
Certain taxa long considered “ancestral reptiles” (e.g., Limnoscelis) have been reinter-
preted as close relatives of Amniota but not members of the crown group (Heaton
1980; Reisz 2007). Furthermore, the phylogenetic analysis by Müller and Reisz (2006)
found Hylonomus and various related taxa to be most closely related to diapsid reptiles,
based on synapomorphies such as the presence of a suborbital foramen in the palate,
the small size of the supratemporal bone, and the long and slender limbs (Heaton and
Reisz 1986).
Tsuji and Müller (2009) defined Eureptilia as the most inclusive clade containing
Captorhinus aguti and Petrolacosaurus kansensis but not Procolophon trigoniceps. Eureptilia
(Fig. 4.1) is distinguished from other amniotes by the absence of a contact between the
postorbital and supratemporal on the skull roof, the narrow blade of the ilium, and the
distinct ventral constriction of the dorsal vertebral centra.

Eureptilia: “Protorothyrididae” + Captorhinidae

Most Pennsylvanian and Permian eureptiles were long placed in two family-level
groups, Protorothyrididae and Captorhinidae (Carroll and Baird 1972; J. Clark and
Carroll 1973).
Protorothyrididae (Romeriidae) was used for various taxa of superficially lizard-like
reptiles, none of which exceeded 20 to 25 cm in total length (Carroll 1969a,b; Carroll
and Baird 1972; J. Clark and Carroll 1973). These forms are known from the Pennsyl-
vanian (Bashkirian) to the early Permian (Cisuralian: Asselian) of North America and
Europe. Hylonomus, from the Pennsylvanian (Bashkirian) of Joggins, Nova Scotia
(Fig. 4.2), is the oldest known reptile. It is documented only by incomplete, disarticu-
lated skeletal remains (Carroll 1964). The supratemporal and tabular bones are small.
Anteriorly, the maxilla bears slightly enlarged teeth; the remaining teeth have small
conical crowns. The vertebrae have narrow rather than “swollen” neural arches, unlike
in other basal reptiles. Müller and Reisz (2006) demonstrated that most “protorothyridids,”
32 The Rise of Reptiles

including Hylonomus and Paleothyris (Figs. 4.2–4.4), are rhinidae as “Anapsida.”) Paleothyris and diapsid reptiles
most closely related to early diapsids. Gauthier et al. share apomorphies such as the long, slender limbs and
(1988b) had already united Paleothyris with Diapsida in a the long, narrow manus and pes (Figs. 4.4, 4.5). The
clade Romeriida. (The original definition of this clade fourth digits of both manus and pes are considerably
did not include turtles, which were grouped with Capto- longer than the other digits, as in many diapsid reptiles.
Sumida (1997) interpreted this feature as indicating ex-
tensive rotation of the hands and feet and push-off in-
volving the elongate digit during locomotion. Thuringo-
thyris, from the early Permian (Artinskian) of Germany
(Boy and Martens 1991), is now considered a stem-
captorhinid, and other “romeriids,” including Romeria,
from the early Permian (Cisuralian: Asselian) of Texas
(J. Clark and Carroll 1973), are basal captorhinids (Mül-
ler and Reisz 2006). Thus, the name “Protorothyrididae”
should no longer be used. Furthermore, the work by
Müller and Reisz and the recent reassessment of the
phylogenetic position of turtles (Chapter 5) have estab-
lished that Williston’s “Anapsida” is not a natural group.
Captorhinidae comprises the most recent common
ancestor of Euconcordia and Moradisaurus and all descen-
dants of that ancestor. It ranged in time from the Penn-
Figure 4.1. Phylogenetic hypothesis of the interrelationships
of Eureptilia. Dots denote node-based clades and parentheses sylvanian (Gzhelian) to the late Permian (Lopingian:
denote stem-based clades. Based mainly on Müller and Reisz Wuchiapingian) and was widely distributed, with rep-
(2006) and Pritchard and Nesbitt (2017). resentatives recorded from the American Southwest,

Figure 4.2. Disarticulated

skeletal remains of the
eureptile Hylonomus lyelli, the
oldest reptile known to date.
Courtesy of Robert Reisz.
 Basal Eureptilia and Diapsida 33

various countries in Europe and Africa, Brazil, China, each maxilla and dentary. The cranial bones have distinc-
and India. Müller and Reisz (2005) found four diag- tive sculpturing (Fig. 4.6). The neural arches are broad
nostic synapomorphies for this clade: reduction in the and swollen, with the zygapophyses positioned far later-
number of maxillary teeth; the anterior position of the ally. The ribcage is broad. The robust, rather short limbs
parietal foramen; the absence of an ectopterygoid; and end in broad hands and feet (Fox and Bowman 1966;
the absence of a tabular. Heaton and Reisz 1980; Fig. 4.5). Captorhinus, from the
Captorhinids include small-sized basal forms with a early Permian (Cisuralian: Asselian-Sakmarian) of Okla-
single row of marginal teeth in each jaw, as well as often homa and Texas and possibly Brazil (Cisneros et al. 2015),
large-bodied derived taxa with multiple rows of teeth in has either a single row or at most three or four rows of
marginal teeth in each maxilla and dentary (Heaton
1979; Kissel et al. 2002). It attained a total length of up to
40 cm. Labidosaurus, from the early Permian (Cisuralian:
Artinskian) of Texas (Modesto et al. 2007; Fig. 4.7), and
Labidosaurikos, from the early Permian (Cisuralian: Kun-
gurian) of Oklahoma and Texas (Dodick and Modesto
1995; Fig. 4.8), have additional rows of teeth lingual to
the marginal tooth rows in each maxilla and dentary. In
captorhinids with multiple tooth rows, new teeth were
added on the lingual side of the jaw and, during growth,
“drifted” toward the labial side of the jaw because of the
addition of new bone lingually and the resorption of
bone labially (de Ricqlès and Bolt 1983). The basal capto-
rhinid Euconcordia, from the Pennsylvanian (Gzhelian)
of Kansas, still replaced its marginal teeth in a manner
similar to that in iguanid lizards, with replacement teeth
erupting directly lingual to the base of the functional
tooth (LeBlanc and Reisz 2015). The pattern of wear on
the teeth and the structure of the jaw joint in captorhi-
nids with multiple tooth rows indicate that the lower
rows bit between the upper ones and that the mandible
was capable of fore-and-aft motion. The largest known
captorhinid, Moradisaurus, from the Permian of Niger,
Figure 4.3. A, skeleton of the basal eureptile Paleothyris attained a skull length of more than 40 cm and has at
acadiana. No scale provided. B, skull of Paleothyris in lateral view. least nine rows of teeth in each maxilla and dentary (de
A, courtesy of the late Don Baird; B, modified from Carroll (2009). Ricqlès and Taquet 1982).

Figure 4.4. Reconstructed

skeleton of Paleothyris
acadiana. Modified from
Carroll (2009).
Figure 4.5. Right pes of A,
Paleothyris acadiana, and B,
Captorhinus laticeps. Abbre-
viations: as, astragalus; ca,
calcaneum; ce, centrale. Arabic
numerals indicate distal carpals
and Roman numerals digits.
Note that the astragalus of
Captorhinus frequently shows
sutures (indicated by dotted
lines) between the original
tarsal elements that became
fused during ontogeny.
Modified from Heaton and Reisz
(1986).

Figure 4.6. A, reconstructed

skeleton of the captorhinid
Captorhinus laticeps in lateral
view. (Distal end of tail
omitted for layout purposes.)
B– C, skull of Captorhinus
laticeps in B, lateral, and C,
palatal views. A, courtesy of
Robert Reisz; B–C, from Heaton
(1979).
 Basal Eureptilia and Diapsida 35

Figure 4.7. Skeleton of the

captorhinid Labidosaurus
hamatus in dorsal view.
Courtesy of Department of
Paleobiology, National Museum
of Natural History.

Figure 4.8. Right Eureptilia: Diapsida

maxillary dentition
By far the most diverse group of present-day land-
of the captorhinid
Labidosaurikos dwelling amniotes, Diapsida includes all extant reptiles
meachami in occlusal and birds. It also has a rich and varied fossil record ex-
view. Courtesy of tending all the way back to the Pennsylvanian and en-
Robert Reisz. compassing an amazing variety of reptiles that lived
on land, in the sea, and in the air. Extant diapsids
belong either to Lepidosauria, comprising the tuatara
(Sphenodon) and squamates (lizards including snakes),
or to Archosauria, comprising crocodylians and birds.
Furthermore, recent phylogenetic studies have shown
that turtles also form part of Diapsida (Chapter 5).
Diapsida comprises the most recent common ances-
tor of Araeoscelidia and Sauria and all descendants of that
ancestor (Laurin 1991). The key diagnostic feature for this
major clade is the presence of upper and lower temporal
openings. The upper temporal (supratemporal) fenestra
is bounded by the parietal, postorbital (and, if pre-
sent, postfrontal), and squamosal. The lower temporal
(infratemporal) fenestra is framed by the jugal, postor-
bital, quadratojugal, and squamosal. The development of
Small-bodied basal captorhinids probably subsisted on these openings results in the formation of two bony
arthropods and other small animals, whereas the larger- arches. This temporal configuration is greatly modified
bodied forms with multiple tooth rows could feed on in many diapsid groups that lack either one (e.g., many
high-fiber plant matter (Dodick and Modesto 1995; squamates) or both (e.g., snakes) of the bony arches. Some
Hotton et al. 1997; Reisz and Sues 2000). diapsid reptiles have only upper temporal openings,
Other documents randomly have
different content
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apercevez pas les suites irréparables. Votre enthousiasme vous
aveugle. Pensez-vous qu'on puisse si facilement braver l'opinion,
qu'on puisse dire impunément: L'opinion est vile, méchante,
déshonnête, je me passerai d'elle pour satisfaire ma conscience et
mon droit? L'opinion se venge, et cela d'autant plus cruellement
qu'on l'a plus justement méprisée. Je la hais comme vous: elle est
perfide et ridicule. Tant qu'on ne l'attaque que par des paroles, elle
ne se formalise pas trop: elle se sent si forte, qu'elle sourit à ses
censeurs, lorsqu'ils l'apostrophent avec esprit ou éloquence. Elle sait
bien que ses plus vifs détracteurs sont les premiers à conformer leur
conduite à ses arrêts. Et c'est là son triomphe. Mais oser lui résister
par ses actes? Oh! c'est terrible. Regardez autour de vous: où sont-
ils les révoltés et les réfractaires? Dispersés, mutilés, anéantis. Eux
aussi étaient braves, croyants, affamés de justice et de bonheur.
Mais ils présumaient trop de leur armure et de leur sainte cause; le
monstre les a étreints et broyés.
Pauline écoutait avec impatience. Pour la première fois, il lui
arriva de s'irriter de ce que lui disait son amant. Une sourde colère
gonflait ses veines. Quoiqu'elle sût bien qu'au fond Odon pensait
exactement comme elle et que, s'il parlait ainsi, c'était moins par
conviction que pour sauvegarder sa responsabilité, elle lui en voulait
de lui répéter ces trop sages raisonnements qu'elle s'était faits elle-
même déjà cent fois.
Elle ne voulait plus discuter. Son parti était pris maintenant.
Revenir en arrière et éterniser d'inutiles débats ne servait qu'à
l'entêter davantage.
Brusquement cruelle, et visant au cœur, elle s'écria:
 —Tu ne m'aimes pas!
Odon pâlit. Il esquissa un geste de supplication; mais il n'eut
pas le temps de prononcer un mot.
—Non, tu ne m'aimes pas, poursuivait-elle avec violence! Si tu
m'aimais vraiment comme je veux qu'on m'aime, tu ne résisterais
pas par de froides raisons à ma volonté faite de passion et de
larmes. Entends-tu? Il n'y a plus place chez moi pour de vaines
controverses. Je souffre trop! Je meurs, si ma vie ne se transforme
pas immédiatement. Aurais-tu peur de me prendre, de m'enlever, de
me soustraire à mon odieuse existence? Oh! je sais que tu ne
m'abandonneras pas, comme le comte des Urgettes a abandonné
Mme de Saint-Géry! Mais peut-être crains-tu le jour où nous
n'aurions plus que nous pour horizon, où nous devrions fuir Paris
pour quelque lointaine campagne, où l'amour serait notre suprême
et universelle ressource. Si tu ne m'aimes pas assez pour me suivre,
je suis perdue. M'aimes-tu, dis-moi? M'aimes-tu?
—Pauline! gémit Odon, entraîné par la passion de sa maîtresse
et comprenant qu'il ne s'agissait plus que de répondre par tout son
amour à l'amour sans bornes dont il se sentait enveloppé. Pauline, tu
doutes de moi!
—Non, non, répliqua-t-elle avec exaltation. Tu es mon ange,
mon salut, mon tout! Mais que suis-je pour toi, moi, femme que tu
aimes, sans doute, que tu n'aimes peut-être pas au point de
consentir joyeusement aux sacrifices qu'exigerait de toi
l'exclusivisme de notre liaison? Car s'aimer, à notre époque inique,
s'aimer c'est se séparer du monde, c'est s'enfermer dans le cloître du
sentiment, c'est perdre son droit à la vie sociale pour conserver son
droit à la vie du cœur. Es-tu prêt comme je suis prête? Si je savais
que tu dusses regretter quelque chose, j'hésiterais, je reculerais: car
plutôt souffrir, plutôt mourir que t'imposer un regret! Parle, dis-moi
franchement si tu m'aimes assez pour qu'à l'idée de me suivre tu ne
sois pas même troublé par l'ombre d'un renoncement.
 —Je t'aime, je ne vois que toi! dit Odon.
—Oh! merci, merci! murmura Pauline de toute son âme.
—Comment pourrais-je ne pas t'aimer assez? T'aimer assez! Il
n'y a pas de degrés dans mon amour: je t'aime. Ce qui n'est pas toi
n'est rien, rien, rien.
—J'en étais certaine, reprit Pauline: je n'ai pas douté de toi un
instant.
—Et puisque tu te donnes, comment ferais-je pour ne pas te
recevoir avec adoration et respect? Je suis ébloui seulement d'un
événement si fabuleux; en face d'une situation si poignante, un
tremblement s'empare de moi; j'ai le vertige à te voir dominer avec
une si superbe audace et une si noble confiance le gouffre
épouvantant de la vie contemporaine. Ah! tu es étrangement belle!
Et malgré que je te connaisse comme la plus remarquable des
femmes, j'ose à peine croire encore à ton incroyable héroïsme.
—Pourquoi nous épuiser à dénouer le nœud gordien, lorsqu'il
est si simple de le trancher?
—Si simple: à condition d'en avoir le courage.
—Ah! mon Odon, s'il ne suffisait que de cela pour conquérir la
vraie liberté! Mais je ne me le dissimule pas: ce ne sera pas la liberté
de l'amour, ce ne sera que la liberté de nous aimer. La vraie liberté
supposerait le consentement unanime des hommes: nous n'aurons
que celui de nos deux consciences, de la nature qui nous bercera et
de Dieu qui nous bénira.
—Ne souhaitons point l'impossible: tenons nos regards fixés sur
la beauté de ce qui est. De par ta volonté, nous sommes libres,
libres de nous aimer. Qu'il nous soit indifférent que les autres
reconnaissent en nous cette liberté! Nous la prenons.
—Et ce n'est point un coup de tête, dit Pauline; j'y ai réfléchi
longtemps; tu as assisté toi-même à la longue et douloureuse
genèse de cet affranchissement. Maintenant que ma décision est
irrévocable, je me sens soulagée du poids terrible qui m'oppressait.
Je suis joyeuse et légère, comme si j'avais à recommencer la vie.
Odon reprit gravement:
—C'est, en effet, une nouvelle vie. Songes-y une fois de plus
avant de creuser entre celle-ci et l'ancienne l'abîme infranchissable.
—L'abîme est déjà creusé. Quoiqu'il ne soit encore visible que
pour moi, il est déjà creusé et déjà infranchissable.
—Tes relations?
—Je les abandonne avec joie au tourbillon des vanités.
—Tes parents?
—Je n'ai plus de parents, sauf ma vieille tante, si affaiblie par
l'âge, si débile d'esprit, qu'elle ne se rend compte de rien. Ma mère
est morte, mon excellente mère... et mon père, mon père si bon, si
touchant... Heureusement qu'ils ne sont plus! Ils n'auraient pas
compris. Si leurs âmes vivent encore, elles savent ce qui est bien.
—Ton mari?
—Lui! c'est surtout lui qui a causé mes souffrances morales. Ai-
je le droit de le tromper, cet homme que je n'aime pas, mais qui n'en
a pas moins reçu de moi le serment de fidélité? A la fois trop
honnête, trop sévère, trop grossier de sentiments et trop imbu de
préjugés, il ne se prêterait pas à ce qu'il appellerait une complicité, il
ne saurait être l'époux complaisant qui, s'apercevant qu'il n'est pas
aimé, tacitement accorde à sa femme la liberté et, au besoin,
favorise son bonheur. Je devrais le tromper, continuer à le tromper,
bassement, perfidement, m'accommoder aux partages et aux
vilenies de l'adultère. Je ne le puis pas, je ne le puis plus. J'ai honte
d'avoir remis jusqu'à présent cette nécessaire purification de ma vie.
Je n'en veux pas à mon mari; il est conséquent avec lui-même: c'est
à moi que j'en veux d'avoir trompé cet homme, qui n'a eu que le
tort, en somme, de ne pas discerner dans la petite fille qu'il a
épousée la future femme passionnée peu propre à goûter les
charmes de l'existence bourgeoise qu'il lui ménageait. Ah! oui, j'ai eu
tous les remords de l'adultère. Mais au lieu de revenir à mon mari,
ce qui serait une tromperie plus abominable encore, je vais à mon
amant.
La vision de ce mari auquel il allait prendre sa femme flotta un
instant dans l'esprit de Rocrange.
«Si c'était à moi qu'un autre enlevât Pauline!» pensa-t-il, sans
pouvoir soutenir plus d'une rapide seconde cette effrayante
hypothèse.
Il savait que Facial n'aimait pas, ne pouvait pas aimer Pauline
comme lui l'aimait. Ne se produirait-il pas, néanmoins, chez ce
malheureux, un déchirement profond, une blessure peut-être
mortelle?
—N'as-tu pas pitié de lui? demanda-t-il.
—Pitié? répondit Pauline en secouant la tête. Son amour-propre
souffrira plus que son cœur. Je n'éprouve pas de réelle pitié pour qui
n'a pas connu le réel amour.
—Que fera-t-il, lorsqu'il apprendra la vérité?
—Rien d'extraordinaire.
—Se battra-t-il?
—Non. Pourquoi? C'est un homme raisonnable. Il réglera
légalement notre situation par le divorce.
—Il ne cherchera pas à te reconquérir en pardonnant?
—Jamais. Ayant violé les lois du mariage, je ne mériterai plus
d'être sa femme. Il me répudiera avec mépris et dignité.
 C'était là, en effet, le vrai Facial: dans les questions de cœur,
moins sujet au désespoir qu'à l'indignation, moins disposé à pleurer
qu'à sévir. Et Rocrange comprit qu'il n'avait que faire de le plaindre.
Toute pitié devait, au contraire, aller à cette pauvre femme, si
sensible, si vibrante, broyée si longtemps dans l'étau du mariage
moderne. Oh! comme elle avait besoin d'être aimée maintenant, et
comme il fallait réparer par une ardeur de baisers et d'adorations le
passé lugubre! Odon entourait sa bien-aimée de ses bras, semblait
la protéger contre l'entreprise inhumaine de la loi, l'arracher aux
étreintes du sort plein de complots. Il contractait avec émotion vis-à-
vis d'elle des devoirs extraordinaires: non pas de ces devoirs factices
et pénibles auxquels obligent la plupart des situations de la vie, mais
de ces devoirs irrésistibles, passionnants, qui ne sont plus même des
devoirs, tellement ils accaparent l'âme. Quelle gratitude emplissait
son cœur! Il éprouvait cette grande volupté de ne pouvoir assez
reconnaître la confiance qui lui était témoignée. Et pourtant, il se
sentait libre. Il était bien entendu entre eux qu'ils s'aimaient
librement, qu'ils se donnaient librement l'un à l'autre, qu'ils restaient
libres jusque dans leurs serments d'amour, si parfois l'entraînement
de la passion les portait à s'en faire. Le jour où ils ne s'aimeraient
plus, si ce jour jamais pouvait luire, ils n'exerceraient l'un sur l'autre
aucune tyrannie. Ils auraient aimé. Ce bonheur leur suffirait. Et il
semblait à Odon qu'à ne pas se lier il en aimait mille fois plus
Pauline. Il eût pris tous les engagements qu'il eût plu à celle-ci de lui
dicter: car l'intérêt de sa maîtresse était la seule chose à quoi il
songeât. Mais elle voulait qu'il n'y eût pas d'autre lien entre eux que
leur amour. Et n'était-ce point leur véritable intérêt à tous deux? Et à
se savoir si libres, ne goûtaient-ils pas davantage le charme d'une
liaison exempte de calculs, où les seules fibres du cœur les
attachaient plus sincèrement que toutes les promesses? Oh! il
l'aimait à tomber à ses genoux, à s'évanouir de joie en sa sainte et
lumineuse présence. Que faisait le mari entre eux deux? Il n'était
bon qu'à être foulé aux pieds, rejeté, expulsé, pour oser mêler
l'arrogance de ses droits caducs à leurs divins épanchements.
 Mais tout à coup une pensée terrible vint bouleverser Odon.
Comment n'avait-il pas réfléchi à cette objection formidable? Et
comment Pauline... Oh! c'était impossible!...
—Ton fils? bégaya-t-il.
Le visage de Pauline ne se troubla pas.
—Ton fils! ton enfant! ton Marcelin pour lequel ton cœur de
mère bat aussi fort que ton cœur d'amante pour moi, l'as-tu donc
oublié? Cette seule apparition ne va-t-elle pas renverser d'un souffle
l'édifice présomptueux de notre amour?
Odon attendait, haletant.
En une appréhension fatale, il eut la vision de l'enfant rappelant
la mère, sinon au devoir, du moins au sacrifice. Il trembla devant la
puissance des bras tendus criant: Ma mère, je suis le lien sacré qui
vous unit indissolublement à mon père! Briserez-vous ce lien? Me
priverez-vous de mon protecteur naturel, de celui qui m'a engendré,
de mon père? Et qui vous dit que je ne l'aime pas, mon père? Est-il
moins mon père que vous n'êtes ma mère? Avez-vous le droit, après
m'avoir mis au monde, en collaboration avec lui, de dissoudre la
famille dont je suis né? L'avez-vous ce droit? Ah! moi, l'enfant, je
suis là, et pour moi vous devez tout supporter, tout souffrir. Il vous
est défendu de changer, par votre bon plaisir, les conditions de ma
naissance. Le sang parle. Le sang est plus fort que tous les caprices;
il prime même les passions les plus irrésistibles et ordonne d'y
résister. Moi, qui suis là, je vous interdis de vous unir à un autre,
tant que mon père est vivant.
Et pourtant, Pauline avait l'air de ne pas entendre cette
supplication filiale.
Que se passait-il dans sa tête qui restait calme, comme si Odon
ne venait pas d'évoquer devant elle le plus redoutable adversaire de
leur amour? Odon considérait sa maîtresse, l'interrogeant du regard
avec anxiété, étonné de ne pas la voir changer de couleur, se
troubler, pleurer, se tordre les mains.
Pauline n'avait pas sourcillé: la question était depuis longtemps
résolue pour elle. Mais elle hésita quelques minutes devant l'aveu
qu'elle avait à faire à son amant.
Ce fut d'une voix très basse, quoique extrêmement tranquille,
qu'elle prononça enfin:
—Mon mari n'est pas le père de mon enfant.
Odon tressaillit. Une sueur froide couvrit subitement ses
tempes.
—Que dis-tu? fit-il, avec effort.
Pauline répéta ce qu'elle venait de dire, mais avec un léger
tremblement, alarmée qu'elle était de l'effet que cette révélation
semblait produire sur Odon.
Rocrange se dressa violemment. Il fit quelques grands pas dans
la chambre, comme frappé de folie, la tête entre les mains et
poussant de rauques exclamations.
—Odon! Odon! gémit Pauline consternée.
Odon s'avança sur elle, lui saisit les poignets et les yeux égarés
cherchant ses yeux pour les fixer furieusement:
—Tu as eu un autre amant que moi? vociféra-t-il... Ah! tu as eu
un autre amant que moi?
Une jalousie atroce le remuait, jalousie brutale, irraisonnée, qui
venait de s'abattre sur lui et de l'étreindre, quoique l'instant
d'auparavant il se fût refusé à croire qu'il pût être sujet à une
pareille passion.
—Réponds! réponds, Pauline! criait-il. Quel est l'homme qui est
le père de ton enfant? Quel est celui qui t'a possédée d'amour avant
moi? Ah! je te croyais pure, et voici que tu as eu un amant, un
amant que tu as aimé comme moi, plus que moi peut-être! Pauline,
tu viens de déchirer mon cœur effroyablement.
Des larmes jaillissaient de ses yeux et devant ce désespoir
Pauline se sentait défaillir.
Mais elle réagit de toute l'énergie dont son âme était capable.
Maîtrisant l'affreuse émotion qui la poignait, elle attendit qu'Odon
eût exhalé le premier flot impétueux de sa douleur; et lorsqu'il se fut
tu, la poitrine seulement secouée encore de sanglots, elle
commença, d'une voix qu'elle fit le plus douce et le plus calme
possible:
—Oui, Odon, j'ai eu un amant avant toi, et si je ne te l'ai pas dit
jusqu'ici, c'est qu'au moment où je t'ai aimé il ne jouait plus aucun
rôle dans la mémoire de mon cœur. J'avais encore moins à te parler
de lui que de mon mari. Il est mort d'ailleurs, cet homme avec qui
j'ai connu les fausses joies de l'adultère, il est mort, et son souvenir
est mort depuis longtemps. Si cet enfant n'était pas là, pour me
rappeler parfois son père, évoquer de l'oubli cette figure disparue,
qui a pu jadis, alors que je n'avais pas accompli le pèlerinage de
l'amour, m'en dresser le fantôme à un coin de ma route, si cet
enfant, qui fait mon orgueil, ne m'inspirait en quelque sorte une
reconnaissance rétrospective pour celui qui me le donna, je n'aurais
qu'un regard d'amertume à jeter sur un passé vide et morne. Je ne
l'ai point aimé, cet homme qui fut mon amant. Mérite-t-il ce titre? Il
n'a su ni dompter mon âme, ni éblouir mes sens. Je suis restée
froide et désolée comme après une effroyable ironie. Pourquoi t'être
livrée à lui? diras-tu. Hélas! c'est pour la même raison qui m'a fait
épouser mon mari. La femme cherche toujours à aimer. Jusqu'au
moment où elle aime vraiment, où elle sait à n'en pas douter qu'elle
aime, bien des tentatives infructueuses ont lieu. Où sont-elles les
privilégiées qui ont trouvé du premier coup l'amant prédestiné et ont
eu l'ineffable gloire de s'offrir vierges à ses baisers? S'il y en a
auxquelles fut départi ce bonheur, qu'elles l'imputent à une faveur
spéciale de la providence. La plupart, j'entends de celles qui aiment,
ont à éprouver l'amère vanité des désirs humains, avant d'en
connaître la possible et magnifique floraison. Heureuses,
bienheureuses encore quand elles la connaissent! O mon Odon,
vierges! Étais-je moins vierge parce que mon corps avait été
possédé? Mais c'est toi, c'est toi qui m'as rendue femme!
Auparavant, quoique femme mariée et femme adultère, je n'étais
pas encore femme. Il me manquait le sens divin de l'amour. C'est toi
qui m'en as dotée: ou plutôt qui l'as découvert, excité, fécondé en
moi. N'as-tu point eu ma vraie virginité? N'es-tu point mon premier,
mon seul, mon parfait amant, mon époux et mon maître? Odon,
Odon, c'est toi que j'aime, je n'ai aimé que toi!
Odon sanglotait toujours, mais son regard s'était adouci. Il
comprenait qu'il avait eu tort de s'emporter et que cette femme
admirable ne perdait en rien de sa valeur pour avoir erré, longtemps
erré à la recherche de l'inappréciable trésor. Lui-même avait eu des
maîtresses, et en grand nombre: et osait-il dire qu'il n'en avait pas
aimé quelques-unes? Et pourtant, lui aussi se sentait vierge, vierge
par le renouvellement qu'apporte tout amour.
—Je ne t'en veux pas, Pauline, prononça-t-il, mais à voix triste
encore.
Il ne pouvait pas se remettre si vite du coup inattendu qui l'avait
frappé, quoique sa raison eût déjà pris le dessus et lui représentât
l'injustice de sa douleur.
Pauline continua:
—Et l'eussé-je aimé, l'eussé-je aimé comme je t'aime, te serait-il
permis de conclure que mon amour actuel n'est pas entier et sans
mélange? Ne devrais-tu pas, au contraire, être fier d'avoir aboli dans
mon cœur les autres sentiments qui auraient pu le partager? Enfin,
et avant tout, n'étais-je pas libre de me donner, alors que je ne te
connaissais pas et que je n'aurais pu me donner à toi? D'où viendrait
que, même dans le cas où j'aurais aimé, tu pusses être peiné de
mon passé?
 —C'est vrai, dit Odon, j'ai agi sous l'empire de la folie:
pardonne-moi.
—Je n'ai rien à pardonner: pour folle qu'elle était, cette jalousie
était de l'amour.
—Pardonne-moi, Pauline, je t'ai offensée. En poussant mon cri
d'indignation égoïste et dément, je me suis ravalé au niveau des
tyrans et des pharisiens, qui entendent bien que la loi soit violée,
mais à leur profit seulement. Le cœur est le cœur: comment
exigerais-je qu'il reste enseveli sous un linceul de mort jusqu'au
moment où j'apparais pour lui souffler la vie? Si ton cœur n'avait pas
été agité depuis longtemps par l'éternel désir, te serait-il possible
maintenant de m'aimer comme tu le fais? Oh non! et j'étais ridicule
de supposer que, douée de passion, tu fusses demeurée jusqu'ici
sans risquer un pas à la poursuite du bonheur. Que tu te sois déjà
donnée, que tu en aies aimé un, deux, plusieurs, qu'ai-je besoin de
m'en préoccuper, aujourd'hui que tu es à moi et que je te tiens
frémissante dans mes bras? Le présent et l'avenir sont la seule chose
qui compte; le passé en a été la préparation; et si le présent
charme, c'est que le passé a été ce qu'il devait être. Pardonne-moi,
Pauline: tu m'aimes, et je ne veux savoir que cela.
La noblesse de ces paroles toucha vivement la jeune femme.
Elle n'était cependant pas entièrement satisfaite: les efforts d'Odon
pour se dompter étaient trop visibles. Elle voulait que son amant
n'eût contre elle pas même l'ombre d'un de ces griefs secrets, dont
on rougit, qu'on est le premier à condamner, mais qui n'en
tourmentent pas moins le cœur.
—Je crains que tu ne m'en veuilles, au fond, dit-elle. Avoue que
j'ai descendu quelques marches du piédestal sur lequel tu te plaisais
à m'ériger.
—Au premier moment, oui, répondit Odon. Je ne réfléchissais
pas que dix ans de mariage avec un mari qu'on n'aime pas justifient
toutes les conséquences.
 —Je n'ai pas besoin d'être justifiée, mais d'être comprise.
Elle lui raconta l'histoire de son adultère. Elle n'en céla ni les
hontes, ni les déboires; elle insista même sur le côté navrant de
cette aventure. Elle se dépeignit telle qu'elle était à cette époque:
irritée de la désillusion de son mariage, impatiente d'aimer, prenant
pour de l'amour les moindres palpitations de son cœur
inexpérimenté, et finalement donnant dans le premier panneau
tendu sous ses pieds par un bel égoïste. Oh! elle n'avait pas été
longue à s'apercevoir de sa bévue; mais elle s'y était entêtée,
espérant toujours, malgré tout, jusqu'au moment où la brutalité
indubitable des faits l'avait laissée gisante sur le carreau, à jamais
rebutée, croyait-elle, de chercher le bonheur par l'amour. Cette
expérience lui avait suffi. Elle avait réfréné en elle ses besoins de vie
sentimentale. Elle en était arrivée à douter de l'amour, ou du moins,
car elle ne le sentait que trop bouillonner stérilement dans son sein,
à douter que sa réalisation fût possible sur la terre.
Odon l'écoutait parler, et, peu à peu, à mesure qu'il pénétrait
mieux le passé de celle qu'il aimait, passé que, quoiqu'il se défendît
de désirer y toucher, elle tenait à lui faire connaître dans ses détails,
le sentiment pénible qui l'avait ému se transformait en ardente
sympathie.
—Pauvre amie! répétait-il, tandis que se succédaient les stations
de ce calvaire.
La pitié gonflait son cœur et n'y laissait plus de place pour la
moindre amertume. Pauline savait si bien le mêler à sa vie, qu'il en
éprouvait lui-même les impressions, la sentait, la comprenait, et
partant n'avait plus rien à en pardonner ou à en excuser. Bien plus, à
voir cette âme se dévoiler davantage, il concevait d'elle une
admiration toujours plus profonde, car il s'étonnait de trouver qu'elle
avait tellement eu soif d'idéal et depuis si longtemps avait souffert
de la disproportion entre ses aspirations merveilleuses et l'indigence
du sort qu'elle avait subi.
 —Et il y a huit ans que cette histoire s'est passée? demanda-t-il,
lorsqu'elle eut fini.
—Il y a huit ans.
—Et depuis?
—Depuis, ce fut la mort de mon âme, ou plutôt, car ses
blessures étaient bien vives, son affreux supplice, l'enfer du doute,
du désespoir, de la fausse résignation, qui cherche à maintenir la
révolte, sans parvenir à autre chose qu'à doter le visage du masque
d'indifférence et de politesse sous lequel les passants ne sauraient
deviner qu'un monde terrible palpite: jusqu'au jour providentiel où je
t'ai rencontré, mon Odon, et où j'ai cru que l'univers allait s'effondrer
sur moi, pour avoir trouvé, enfin! enfin! le bonheur dans deux bras
amis.
—N'as-tu vraiment pas essayé durant ces huit ans de te donner
à un autre homme?
—Non, fit Pauline: l'amour que je concevais était si haut, qu'il
me semblait impossible qu'il se trouvât quelqu'un capable d'y
répondre. Bien des hommes m'ont fait la cour; en tous je démêlais
l'égoïsme cynique, la sensualité grossière, la vanité stupide. Aucun
ne m'aimait vraiment, et, comme avec les années l'idéal que je me
créais de l'amant se complétait et grandissait, aucun, même parmi
les meilleurs, ne me paraissait digne d'être aimé. Au spectacle des
misérables intrigues qui se nouaient et se dénouaient autour de moi,
je n'étais que plus décidée à abandonner aux âmes médiocres de si
méprisables commerces. J'avais renoncé à croire; la foi était partie
enlevée par les serres de la déception. Il fallait un miracle pour me
sauver: le miracle s'est produit. Dieu que j'avais renié s'est manifesté
au moment où je ne m'attendais plus qu'au néant, et je suis
maintenant en adoration devant sa bonté et sa puissance.
—O Pauline! dit Odon, tu es la plus noble, la plus rare des
créatures. Je suis un misérable de t'avoir soupçonnée d'une
faiblesse. Une faiblesse, bon Dieu! Quelle prétention avais-je? Mais
je te voulais sans tache, comme la divinité pure à laquelle on a
dressé un autel et qu'on pare de toutes les vertus. Et, mauvais
croyant, il m'avait semblé qu'un nuage passait sur ta blancheur
immaculée. Mais, voilà que tu m'apparais maintenant plus
éblouissante qu'avant. Oh! pardonne, pardonne!
Cette fois, c'était sincère et profond. Ce n'était plus seulement
sa raison qui le poussait à rendre justice, mais tout son cœur.
Les yeux de Pauline brillèrent de joie, son âme rayonna.
Odon s'était agenouillé devant elle. Il baisait les plis de sa robe;
et sur sa main, la jeune femme sentit tomber une larme.
Ce fut un instant de muette extase. Puis, lorsqu'il se fut relevé,
elle se jeta dans ses bras, comme pour y chercher la protection
suprême.
—Rien ne pourra m'arracher de toi! balbutiait-elle.
—O mon amie, je serai ton seul, ton véritable époux. Je le vois
maintenant, le monde ne saurait être pour toi qu'un désert; la
famille même, cette prison où tant, qui soupirent après la liberté,
sont retenus par de multiples chaînes, est démolie autour de toi et
ne t'offre que des ruines inhabitables; tout t'éloigne de celui auquel
la loi t'a lié, tout et jusqu'à l'enfant, qui d'habitude est l'inexorable
carcan rivant au même collier de fer deux têtes ennemies. Je n'ai
plus d'objection, plus. Je suis convaincu que ton bien comme ton
devoir consistent à abandonner ton mari pour me suivre. Je
n'appréhende plus pour toi ni les regrets, ni les défaillances. Au point
où tu en es, la seule solution possible, c'est la rupture avec un passé
de larmes et de mensonge.
—L'honneur même, cet honneur dont on a plein la bouche et
qu'on comprend si peu, l'honneur même l'exige.
—Je ne te parle pas de ma joie, Pauline; elle est immense. Oh!
nous serons heureux!
 —Je le veux, Odon.
—Un avenir de bonheur caché, loin de la foule, loin des vanités
et des perfidies, s'ouvre devant nous. Une idéale confiance en Dieu,
en la justice, en l'amour remplit nos âmes. Unis par le saint mystère
d'une même foi, nous oublierons les hommes, les païens, les
barbares. Nous les laisserons à leurs faux dieux et à leurs cultes
malfaisants. Chère épouse, tes yeux seront mon univers, tes beaux
yeux où se révèle l'unique grâce qui me touche. Peu nous importe le
bruit que l'on fera sur nous: il ne parviendra point à nos oreilles.
Nous aurons le témoignage de notre conscience, le seul bien
nécessaire, et qui ne nous faillira pas.
—Oh! oui, dit Pauline, la conscience, l'honnêteté, l'amour!
Elle appuya sa tête sur le sein de son amant.
Une bénédiction semblait planer sur eux. La douceur de cette
heure était si grande, qu'ils ne savaient comment s'exprimer
mutuellement leur gratitude.
Ils restèrent longtemps silencieux en une étreinte bienheureuse.
Puis, Pauline dit:
—Dès demain, mon mari saura tout.
Elle avait à peine prononcé ces mots, qu'un bruit de pas se fit
entendre dans le salon voisin.
Pauline pâlit affreusement.
La portière s'écarta. Sur le seuil de leur chambre, un homme
apparut:
Facial.
 XII

Depuis plusieurs heures, Facial se promenait dans son cabinet,

en attendant l'entrevue qu'il devait avoir avec sa femme.
Un domestique vint lui annoncer que madame était arrivée.
Il se recommanda encore la plus glaciale, la plus dédaigneuse
politesse, boutonna sa redingote, but un petit verre de cognac, et
passa au salon où l'attendait Pauline.
Elle se leva à son entrée et lui tendit la main sans affectation.
—Nous ne sommes coupables ni l'un ni l'autre, dit-elle;
épargnons-nous mutuellement les reproches et les grands mots.
Facial resta abasourdi de ce début. Il se préparait à subir des
attendrissements, des sanglots, une femme se jetant à ses pieds et
demandant grâce, et voici qu'il la trouvait aussi calme que lui.
—Asseyez-vous, Madame, dit-il avec un geste vague.
Ils prirent place en face l'un de l'autre, séparés par une petite
table.
—Je n'ai pas d'explication à vous donner, fit Pauline au bout
d'un instant de silence, et je vous prie de ne pas en exiger de moi. Il
doit vous être assez indifférent de savoir pourquoi et comment j'en
suis venue à rompre les liens qui nous unissaient. Il est probable
d'ailleurs que si je tentais de vous l'expliquer, vous ne me
comprendriez pas. Veuillez donc ne considérer que les faits. Ils sont
trop évidents pour que je songe à les nier ou à les atténuer. J'en
assume la responsabilité.
Facial perdait pied. Il ne concevait pas que Pauline osât se
présenter à lui autrement qu'en pécheresse repentante et accablée
de honte.
—Ah! misérable femme! s'écria-t-il, oubliant d'un coup ses
projets d'impassibilité.
—Ne le prenez pas sur ce ton, dit Pauline, je vous en supplie.
—Comment! Vous m'avez trompé, trahi, déshonoré, vous avez
commis un crime épouvantable, vous voilà souillée, couverte de
boue, et vous venez tranquillement m'annoncer que vous en
assumez la responsabilité! Je crois bien que vous en assumez une de
responsabilité, et effroyable! Les conséquences de votre faute seront
terribles, terribles...
—Il est inutile de vous emporter: ce qui est fait est fait, et si
c'était à refaire, je le referais. Veuillez me dire maintenant quelles
sont vos intentions.
Facial la regardait effaré.
—Mes intentions? mes intentions? Vous en parlez avec une
légèreté... Ah ça! éclata-t-il, pensez-vous que je vais passer l'éponge
sur vos déportements, vous ouvrir de nouveau, comme si de rien
n'était, ma maison et mes bras, vous supplier peut-être—telle est
votre audace!—de reprendre la vie commune agrémentée de toutes
les complaisances? Ne vous bercez pas d'illusions. Ne vous figurez
pas que votre pouvoir sur moi soit si grand, qu'il vous suffise de
paraître pour reconquérir votre place au foyer. Vous vous traîneriez à
mes genoux, que je resterais inflexible. Madame, je ne suis pas de
ceux qui pardonnent.
Cette phraséologie mettait Pauline au supplice.
 —Je ne suis point venue ici mendier votre pardon, dit-elle. Je ne
saurais qu'en faire. Dites-vous bien d'ailleurs que si vous souffrez
maintenant à cause de moi, j'ai souffert, moi, pendant dix ans à
cause de vous, et ne vous posez pas en accusateur: ce rôle vous
convient peu.
—Quelle impudence! fit Facial avec indignation. Mais vous êtes
un serpent que j'ai réchauffé dans mon sein!
Pauline haussa les épaules.
«Rien, pas un cri du cœur ne lui échappe!» pensait-elle.
Elle se taisait, hautaine, sous les injures que Facial déversait.
Qu'aurait-elle dit? Elle ne pouvait pas lui prêter son cerveau, pour
qu'il sentît avec ses sentiments et comprît qu'il n'avait pas le droit de
la juger. Il voyait à son point de vue, un point de vue abominable et
faux, mais qui était le sien. Que servait alors de répondre?
En proie à une fureur qu'il ne cherchait plus à contenir, Facial se
répandait en discours diffus, boursouflés, pleins de périodes
déclamatoires et d'imprécations violentes. Il dépassait les bornes,
traitait sa femme de fille perdue, la ravalait au-dessous des
prostituées, qui, elles, n'ont juré fidélité à personne. Les outrages
jaillissaient de ses lèvres. Lui, si châtié d'habitude dans son langage,
trouvait d'ignobles insultes à lancer comme des crachats au visage
de celle qui lui était intellectuellement et moralement si supérieure.
Elle ne bronchait pas; pâle, les traits immobiles, elle laissait passer
ce flot d'ordure qui ne l'atteignait pas.
Épuisé, Facial s'arrêta et s'affaissa dans un fauteuil.
—Avez-vous fini? demanda Pauline.
Il se redressa, comme sous un coup de fouet.
—Je n'ai pas encore dit le plus important, Madame, reprit-il
foudroyant; je n'ai pas encore prononcé le mot fatal...
 —Prononcez-le, interrompit-elle, je n'attends que cela.
—Vraiment, Madame, le divorce ne vous fait pas peur?
Il espérait la voir s'abattre sous l'épouvante de ce mot et
mesurer enfin l'horreur de son crime à la grandeur de la punition.
Mais elle ne parut pas s'en émouvoir.
Il accentua d'une voix sévère:
—Le divorce, Madame! le divorce!
—Je suis heureuse, répondit simplement Pauline, que vous
compreniez comme moi qu'une séparation est nécessaire. Vous la
voulez légale, tant mieux: l'ordre est une excellente chose, et ma
liberté en sera moins précaire. Le divorce est la meilleure solution à
notre situation. Si vous avez cru que je me ferais des illusions sur
votre tendresse à mon égard, vos paroles me montrent que vous en
entretenez sur celle que je vous porte. Vous vous imaginez que «ma
faute»—je conserve à mon acte ce nom, puisqu'il est consacré,
quoique ma vraie faute, faute bien involontaire et toute d'ignorance,
ait été de vous épouser sans savoir ce que c'est que l'amour—vous
vous imaginez que ma faute est le résultat d'un de ces coups de tête
ou de sang familiers aux femmes peu scrupuleuses, qui durent le
temps d'un caprice et dont elles se mordent amèrement les doigts,
si, par malchance, le mari découvre et sévit. Vous supposiez que ce
mot de divorce allait me prosterner à vos pieds humiliée et brisée,
pleurant des serments de repentirs éternels. Vous vous trompez. Ma
faute a été voulue et longuement méditée. Bien loin d'en redouter
les conséquences, j'étais à la veille de vous découvrir moi-même la
vérité. Vous m'avez prévenue: ce n'est pas une raison pour que je
change de contenance. Non, je ne crains pas le divorce; je l'appelle,
je le désire. Mais ici vous êtes le maître, vous seul avez qualité pour
le réclamer, puisque, au point de vue de la loi, c'est vous qui êtes
l'offensé.
—C'est bien, Madame, nous divorcerons. Telle était mon
intention: vos bravades ne font que m'y affermir.
 —Sur quoi baserez-vous votre demande?
—Sur la vérité: votre adultère. Songeriez-vous à le nier?
—Oh non, je vous aiderai même à l'établir.
—Il y a des maris chevaleresques qui en pareille circonstance
poussent l'abnégation jusqu'à prendre la faute sur eux. N'attendez
pas de moi une telle délicatesse. Je considère l'adultère, même
l'adultère de l'homme, comme une chose trop grave pour que je
consente à m'en charger. Que m'importe votre honneur, maintenant
que vous l'avez perdu. Le divorce sera prononcé contre vous.
—J'entends. Vous m'offririez d'ailleurs ce petit sacrifice, que je
n'accepterais pas.
—Tout ce que je puis faire, c'est de ne pas vous traîner devant
le tribunal correctionnel pour obtenir votre condamnation. Je
délaisse cette vengeance.
—Quelle magnanimité!
—Le nom de votre complice ne sera pas même prononcé dans
les considérants. Vous pourrez l'épouser, puisque vous prétendez
l'aimer, et essayer de racheter avec lui les torts que vous avez eus
avec moi.
Facial se croyait sublime.
—Il est marié, dit Pauline.
—Il peut divorcer.
—Il ne le peut pas: sa femme est catholique.
Facial leva les yeux au ciel.
—Dans quel abîme êtes-vous tombée! Enfin s'écria-il, vous
l'avez voulu, Madame, vous l'avez voulu!
 —C'est bien. Ne parlons pas de moi. Puis-je vous demander
quelles sont les preuves que vous produirez devant les magistrats?
—Des preuves? J'ai des témoignages, des présomptions
morales, des faits matériels qui, réunis, formeront un dossier
suffisant pour vous confondre.
—Croyez-moi, laissez de côté tout cet arsenal. Il est inutile,
puisque j'avoue. Ne désirez-vous pas, comme moi, aboutir par les
moyens les plus rapides et les plus simples?
—Sans doute, et si vous avouez cela ira tout seul. Mais il faut un
aveu écrit.
—Qu'à cela ne tienne, je vais vous écrire une lettre où je
reconnaîtrai explicitement ma culpabilité.
—Comme vous voudrez, fit Facial. D'habitude, les femmes
n'avouent pas ces choses-là; leur pudeur les pousse à se défendre
même contre l'évidence. Il faut que vous ayez perdu tout sens
moral.
Sans répondre, Pauline ouvrit un buvard, prit une feuille de
papier et écrivit une demi-page qu'elle signa.
—Cela suffit-il? demanda-t-elle en tendant la pièce à son mari.
Facial la lut deux ou trois fois attentivement.
—Cela suffit, dit-il.
Puis il la serra avec soin dans son portefeuille.
—Et maintenant, Madame, termina-t-il, nous ne nous
retrouverons que devant les juges. Que Dieu vous pardonne!
Mais au lieu de partir, Pauline se dirigea vers une porte menant
dans les appartements intérieurs.
—Où allez-vous! cria Facial.
 —Mon fils... Je vais chercher mon fils.
—Pour quoi faire?
—Pour l'emmener.
Il se précipita et lui barra le passage.
—Vous ne passerez pas!
—Monsieur!
—Je vous le défends!
Elle s'arrêta haletante. Un éclair flamba dans ses yeux.
—Vous oseriez me défendre de prendre mon fils? prononça-t-
elle les dents serrées.
—Parfaitement.
—Mais c'est mon fils! rugit-elle.
—C'est aussi le mien, dit Facial.
Une horrible lueur palpita dans l'esprit de Pauline. Son fils! son
fils! Facial songeait à le lui enlever! Oh! c'était impossible! Quelle
monstrueuse pensée venait de germer là tout à coup, si
monstrueuse que pas un instant le soupçon que cela pût se produire
ne lui était venu! La séparer de son fils! Ce forfait épouvantable
serait-il permis? Non, non, elle se trompait, elle avait mal entendu!
Son mari était un homme après tout: il n'allait pas voler un enfant à
sa mère!
—Je veux mon fils! supplia-t-elle la tête pleine de vertige.
—Vous ne l'aurez pas.
Alors, en une abondance éperdue de paroles incohérentes,
pleurant, défaillant, les mains frissonnantes, elle divagua:
 —Vous n'avez pas formé l'infernal projet de m'arracher mon
enfant! Ce n'est pas sérieux, ce n'est qu'une effroyable plaisanterie!
Dites, dites que vous n'avez voulu que me faire peur! Je suis mère,
moi, savez-vous bien? Ce serait me tuer que de m'ôter l'enfant que
j'ai porté dans mon sein, que j'ai nourri, que j'ai élevé, qui est mon
sang et ma vie! Oh! vous savez cela! Vous ne voudrez pas
commettre un crime si infâme! Si vous avez jamais eu pour moi un
sentiment qui ne fût pas de la haine, vous épargnerez la
malheureuse qui a été votre femme, vous n'exercerez pas sur elle
une atroce, une basse vengeance. Vous ne dites rien; vous attendez
que je me sois mieux humiliée. Parlez, que dois-je faire pour vous
fléchir? Oh! grâce! grâce! L'angoisse m'étreint à la gorge, ma voix se
perd, les mots manquent à mon cœur...
C'était enfin la scène que Facial attendait et à laquelle il s'était
préparé. Seulement, au lieu que ce fût la femme, c'était la mère qui
criait grâce.
Il répondit durement:
—C'est trop tard: il fallait songer à cela avant.
Une nouvelle énergie galvanisa Pauline:
—Vous avez l'audace de séquestrer Marcelin? proféra-t-elle avec
un tel emportement, que Facial crut qu'elle allait se jeter sur lui.
—Sa place n'est pas avec vous. Je le garde.
—De quel droit?
—De quel droit? Je crois, Madame, que vous vous méprenez ici
étrangement sur vos droits. Apprenez donc que, le divorce étant
prononcé contre vous, c'est à moi, en principe, que le tribunal doit
confier l'enfant. Il suffit que j'en fasse l'objet d'une demande, et
c'est ce qui sera, pour que, malgré tout ce que vous pourrez arguer,
le droit de garder Marcelin me soit acquis.
 A ces paroles qui éclairaient tragiquement la situation, Pauline
sentit tout s'effondrer en elle.
Un dernier espoir restait, auquel elle s'accrocha désespérément.
Il fallait pour cela l'aveu terrible. Mais plus rien ne lui coûtait.
Se campant devant son mari, le fixant les yeux dans les yeux,
elle dit avec un cinglement:
—Cet enfant n'est pas de vous.
Facial sursauta.
—Il n'est pas de vous, reprit-elle plus ardemment, il est de
M. de Hartwald. Car je vous ai trompé autrefois avec M. de
Hartwald. C'était à l'époque où il était secrétaire d'ambassade à
Paris. Vous vous le rappelez? J'ai fait sa connaissance dans un bal. Il
venait souvent ici. Vous l'invitiez. Eh bien, je vous trompais avec lui.
Pendant un an, je vous ai trompé; et vous ne vous en doutiez pas.
Marcelin est né de cet adultère. Regardez-le, il n'a rien de vous: il ne
vous ressemble ni au physique ni au moral. Remarquez son nez, son
nez droit, fin, distingué, et ses cheveux, ses cheveux blonds: c'est le
nez et les cheveux de M. de Hartwald. Il a, par contre, mes yeux et
ma bouche. C'est frappant. M. de Hartwald est mort; cet enfant est à
moi seule...
Elle s'arrêta, regardant toujours son mari. Mais celui-ci, après un
premier choc de surprise, avait eu le temps de se remettre.
—Ah! par exemple! s'écria-t-il en riant insolemment, vous avez
de l'imagination! Ma parole, à vous entendre, on dirait que c'est
arrivé! Mais ça ne prend pas! Ça ne prend pas! Marcelin le fils de
M. de Hartwald! Elle est bien bonne!
—Vous ne me croyez pas? fit Pauline bouleversée.
—Vous croire? Ah ça, pour qui me prenez-vous? Il est visible
que vous venez d'inventer cette histoire de toutes pièces. C'est un
mensonge, et qui plus est un mensonge ignoble. Ah! Madame, vous
étiez déjà bien bas dans mon estime: vous voici dans la fange
jusqu'au cou.
—Vous ne me croyez pas? répéta-t-elle avec accablement.
—Inventez autre chose, ou mieux n'inventez rien du tout. Votre
paroxysme vous égare jusque dans le ridicule. Marcelin ne serait pas
mon fils! Vous moquez-vous? Vous trouvez qu'il ne me ressemble
pas? Vous êtes donc aveugle! Et la voix du sang, Madame, la voix du
sang! Est-ce que je me sentirais son père, si je ne l'étais pas?
—Mon Dieu! mon Dieu! gémissait Pauline.
Et elle demeurait stupide devant son impuissance à établir la
vérité. Elle ne possédait aucune preuve de ses relations avec M. de
Hartwald. Tout avait été détruit. Il n'existait pas un mot de billet, pas
une photographie, pas un signe, pas un document quelconque, rien,
rien, rien, que sa parole à elle et cette ressemblance qu'elle était la
seule à apercevoir.
Alors, folle, elle cria à son mari:
—Rendez-moi la lettre!
—La lettre?
—Oui, la lettre que je viens d'écrire et où je me reconnais
coupable. Je ne divorce plus.
—Pardon, Madame: vous ne divorcez plus, mais moi je divorce.
Je ne vous rendrai pas la pièce que vous m'avez si légèrement
fournie.
—Oh!...
—D'ailleurs, cela ne vous avancerait pas à grand chose. Comme
je vous l'ai dit, j'ai des témoignages à faire valoir. La procédure sera
un peu plus longue, voilà tout.
 —Je me défendrai, je lutterai et peut-être parviendrai-je à jeter
quelque doute dans l'esprit des juges. Rendez-moi ma lettre!
—Non.
—C'est une lâcheté!
—Une prudence.
—Mon enfant! mon enfant!
Elle voulut s'élancer. Facial la saisit violemment par les bras et la
coucha de force dans un fauteuil. Sans cesser de la maintenir, il
appela:
—Victor!
Le valet de chambre parut.
—Prévenez miss Dobby qu'elle ait à emmener immédiatement
mon fils là où elle sait. Accompagnez-les.
En proie à une indicible horreur, Pauline se débattit
convulsivement. On enlevait son enfant! Elle ne le verrait plus,
plus... C'était fini!
—Le voir, râla-t-elle... je veux le voir...
Mais les deux mains atroces de son mari la serraient comme
dans un étau, la clouaient, la paralysaient.
—Lâchez-moi!... Oh! ayez pitié, pitié!... Mon Dieu, ayez pitié!...
On entendit, du côté de l'antichambre, une lointaine voix
d'enfant:
—Maman! maman!
Pauline se raidit en un suprême effort. Mais ce fut en vain. Elle
retomba brisée sous la masse vigoureuse qui pesait sur elle.
Elle cria.
 Facial lui mit son genou sur la bouche.
Quelques instants épouvantables se passèrent, pendant lesquels
elle crut mourir, tout son pauvre corps tordu comme dans les
spasmes d'une torture.
Enfin, Facial la lâcha.
—Vous êtes libre, dit-il.
Elle se leva d'un bond fiévreux et se précipita à travers
l'appartement. Elle en parcourut hâtivement les diverses pièces. Le
vide, le vide partout. Marcelin n'était plus là. Dans la salle d'étude,
un désarroi de livres et de cahiers... Elle baisa en sanglotant ces
objets que son enfant maniait encore quelques minutes auparavant,
elle les baisa comme des reliques sacrées, et son cœur de mère
éclatait dans sa poitrine... Ses lèvres battaient, ses paupières
tremblaient nerveusement; elle répétait le nom chéri, tantôt tout
bas, comme une prière, tantôt en appels désespérés écorchant sa
gorge en feu. Elle reprit deux ou trois fois sa promenade errante de
chambre en chambre, lentement maintenant, anéantie, s'arrêtant à
chaque détail qui lui évoquait Marcelin. Lorsqu'elle revint au salon,
où Facial attendait qu'elle se fût convaincue de l'inutilité de sa
révolte, elle n'avait plus l'air que d'un spectre désolé, d'une statue
vivante de l'effroi.
La vue de son mari sembla la glacer d'épouvante. Elle porta ses
mains en avant, dans un long geste de répulsion. Quelques mots
rauques sortirent péniblement de sa bouche contractée.
—C'est vous... c'est vous...
Et elle s'abîma sur le tapis, sans connaissance.
Facial sonna la femme de chambre. Il lui montra le corps
inanimé de Pauline. Puis, il prit son chapeau et partit.
Au bout d'une demi-heure, Pauline revint à elle. La femme de
chambre l'avait portée sur un lit, lui faisait respirer des sels,
étanchait avec un mouchoir imbibé d'eau le sang d'une petite plaie
qu'elle s'était faite en tombant.
—Où est mon fils?
—Je ne sais. Il est sorti avec sa gouvernante et Victor.
—Et monsieur?
—Il est sorti aussi. Il n'y a personne à la maison.
Elle s'élança à bas du lit, sans prendre garde qu'elle pouvait à
peine se tenir debout.
—Madame n'est pas encore remise; Madame ferait mieux de
rester couchée.
—Laissez-moi!...
Elle descendit dans la rue, échevelée, hagarde, semblable à une
aliénée.
 XIII

—Que vous êtes agaçant, dit Julienne, on ne peut rien tirer de

vous!
—Mais, Madame, répliqua Réderic, vous m'interrogez à tort et à
travers, vous et ces dames, sur ce que vous vous plaisez à appeler
les mystères de l'affaire Rocrange! Que voulez-vous que je vous
dise? C'est très simple. M. de Rocrange aimait Mme Facial; Mme
Facial aimait M. de Rocrange; Mme Facial, qui, paraît-il, est une
femme sincère, ne s'en est point trop cachée; et M. Facial, qui
n'entend pas plaisanterie, plaide aujourd'hui même en divorce contre
elle. Quoi de plus clair, de plus net, de plus logique? Il n'y a pas
ombre de mystère. Les dessous n'existent pas. Tout cela est
purement honnête.
—Honnête! s'exclamèrent avec des mines effarouchées la
baronne Citre, Mme Sermais et Mme d'Orgely.
—Qu'appelez-vous l'honnêteté? demanda Réderic.
Cette question déconcerta.
—L'honnêteté, c'est de rester fidèle à son mari, risqua enfin la
baronne.
—Oh! ma chère, que vous êtes vieux jeu! ne put retenir
Julienne.
 —En effet, Madame, dit Réderic, c'est là une honnêteté
antédiluvienne.
—L'honnêteté est au moins la bienséance, corrigea la baronne,
consciente d'avoir émis une niaiserie.
—C'est ça, c'est ça! zézaya Mme d'Orgely sous son éventail.
—Et la bienséance? continua Réderic imperturbable.
Cette fois, personne ne hasarda de réponse.
—La bienséance, reprit-il, voici: tromper son mari avec
discrétion et rouerie; s'évader sans bruit de sa tutelle; prendre
subrepticement tout le champ possible pour ses ébats et savoir
revenir en hâte au moindre signal de la laisse, que l'on a tendue
juste à point pour qu'une malencontreuse secousse n'avertisse pas
de l'incartade le légitime propriétaire. Certaines femmes sont tenues
très court; d'autres ont la laisse étonnamment longue: toutes
jouissent autour du poteau marital d'un espace plus ou moins grand
où brouter le thym d'amour. Ah! chèvres bienséantes, au poil blanc,
à l'œil innocent, jouez tant qu'il vous plaît entre les rocs qui vous
dissimulent, derrière les hautes herbes, à couvert des ondulations de
terrain; mais ne vous avisez pas de ronger de vos dents fines la
corde qui vous retient pour aller gambader à l'aise sur les hauts
sommets, où l'air est pur et léger, sans doute, mais où vous ne
seriez plus que de vilaines chèvres sauvages indignes de
considération. Vous aimez la liberté, mais il vous faut une liberté qui
ait l'air de ne pas trop frauder l'esclavage. Vous ne la prenez pas,
vous la dérobez. Vous ne sauriez avoir de désirs vifs, francs, joyeux;
vous ne connaissez que les envies louches, inavouées, satisfaites en
secret comme des vices. L'intrigue est, du reste, votre plaisir. Vous
ne trouveriez guère de charme à l'amour, s'il n'était avant tout le
fruit défendu, auquel il s'agit de goûter par une adroite et perfide
maraude. Vous craignez la passion et vous la haïssez: et lorsque, par
miracle il s'en trouve une qui soit autre chose qu'une coquette ou
une coquine, vous le lui faites expier avec acharnement. Ah! elle ne
trompe pas comme vous: haro sur elle! N'est-ce pas, mesdames, la
bienséance consiste dans la déloyauté d'abord, et dans la cruauté
ensuite?
Réderic avait fait cette petite exécution sur un ton de persiflage
mi-plaisant, mi-acerbe, dont il n'y avait pas lieu de s'offenser, mais
qui n'en était pas moins mordant.
—Voyons, Réderic, fit Julienne assez vexée, vous êtes
insupportable! En avez-vous encore pour longtemps à faire votre
Alceste?
—J'ai fini, belle dame, j'ai fini: le métier est trop peu profitable,
et il vaut mieux hurler avec les loups.
—Le monde est tel qu'il est, et ce n'est pas vous qui le
changerez. Alors?
—Alors, je n'essaye point de le changer. Je constate les petites
crapuleries qui s'y passent, et bien que je ne prenne pas à ces
observations un très vif plaisir, je ne suis pas Alceste au point de
m'en irriter plus que de raison.
—Et vous consentez parfois à hurler avec les loups, suivant
votre exquise expression. Mais, à ce propos, revenons à nos
moutons.
—Les avons-nous quittés?
—Réderic, si vous continuez, je me fâche.
—Ma chère, il veut défendre cette pauvre Pauline et son ami
M. de Rocrange, dit cauteleusement Mme Sermais. Il est charitable
sous son pessimisme. Seulement il procède d'une façon peu
intelligente. Ce n'est pas en s'en prenant aux honnêtes femmes
qu'on reconstituera l'honneur de celles qui s'exposent. Qu'on sollicite
notre indulgence, rien de mieux; nous sommes prêtes à l'accorder;
nous vivons à une époque où l'on est indulgent. Mais que l'on exige
notre respect pour des femmes si peu soucieuses des mœurs
qu'elles semblent trouver du plaisir à se compromettre, c'est
vraiment se moquer de nous.
—Très bien, approuva la baronne.
—Je vois que mes clients, puisque clients il y a, sont bien
malades, fit Réderic sans s'émouvoir. Il ne me reste qu'à les
abandonner à l'inclémence du tribunal.
—Épousera-t-elle au moins son Don Juan? demanda Mme
Sermais.
—Mais, ma chère, dit en riant Julienne, ne savez-vous pas qu'il
existe déjà une Mme de Rocrange?
—Dans quel bourbier pataugeons-nous! déclama la Sénéchale,
qui se délectait à suivre cette conversation.
—Je me le demande, observa Réderic sentencieux.
Julienne se leva et alla lui donner une tape sur les doigts.
—Réderic, je vous intime l'ordre de vous taire. Lorsqu'on vous
interroge, vous vous dérobez, et quand on ne désire plus rien de
vous, vous manifestez votre vilain caractère par de désobligeantes
remarques qui sont peu d'un galant homme.
—C'est dommage que notre incomparable sénateur ne soit pas
là, il ferait mieux notre affaire.
—Ne vous désolez pas, il va venir.
—Vous savez, ma belle, dit la Sénéchale à Julienne, que c'est
exprès pour vous que ce cher homme assiste à l'audience. Il est si
peu curieux de sa nature, et ce linge est si sale à voir laver!
—Ah! fit Réderic, Sénéchal est au Palais?
—Oui, dit Julienne, et nous allons avoir des détails tout frais.
—Quel bonheur! s'écria étourdiment Mme d'Orgely.
 —Il est charmant! soupira la baronne.
—Comme le vicomte et la vicomtesse doivent être ennuyés de
cette aventure, émit la Sénéchale avec componction. M. de Rocrange
s'est comporté...
—Oh! Madame, interrompit Mme Sermais, il a fait son métier
d'homme. Il n'y a rien à lui reprocher. Pour Pauline, quelque pitié
qu'on ait pour elle, il faut avouer qu'elle est coupable. Je dis
coupable plus que malheureuse, car tout dans sa conduite prouve
qu'elle a visé au scandale. Ne lui eût-il pas été facile, même en
supposant le pire, de s'arranger à étouffer l'affaire, à éviter l'odieux
d'un procès en divorce? Mais non, elle a été cassante, elle a rendu la
conciliation impossible. Ce n'est point contre son mari qu'elle est
partie en guerre, c'est contre la société, contre l'ordre, contre nous.
—Cela se pardonne moins aisément, dit Réderic.
—Et maintenant, demanda la baronne, que va-t-elle faire?
—Elle ne peut pas continuer à habiter Paris, dit Mme Sermais.
Personne ne l'a revue, du reste. Pas même vous, chère madame?
ajouta-t-elle en se tournant vers Julienne. Vous étiez pourtant de
son intimité, je crois?
—Moi? pas du tout. Nous nous fréquentions seulement, ou
plutôt elle me fréquentait. Ces derniers mois, je l'avais presque
perdue de vue.
Une pendule se mit à sonner.
—Il devrait y avoir un coq sur cette pendule, dit Réderic.
Une rougeur fugitive passa sur le visage de Julienne. Elle reprit
vivement sans paraître avoir remarqué l'interruption:
—Sénéchal, qui sait tout, m'a affirmé que Pauline était à Grasse.
Aussitôt après l'éclat, elle se serait retirée chez sa tante, puis,
quelques jours plus tard, serait partie pour le Midi. Je suppose
qu'elle est revenue pour le procès, mais je ne saurais vous le dire au
juste.
—Et M. de Rocrange?
—M. de Rocrange est aussi parti.
—Pour le Midi?
—C'est vraisemblable. Réderic pourrait nous renseigner, mais il
ne le fera pas.
—Pourquoi ne le ferais-je pas? Vous voulez savoir où est
Rocrange? C'est bien simple: il est à Béthanie.
—Comment?
—A Béthanie, loin de l'œil des pharisiens, avec Marie, Marthe et
Lazare, fondus pour lui en une seule personne: Lazare qu'il a
ressuscité, Marie et Marthe qui l'aiment, l'une mystiquement, l'autre
candidement.
—Et pendant ce temps, dit Julienne avec un haussement
d'épaules blagueur, on conspire contre lui dans le Sanhédrin! Pour
Dieu, Réderic, mon pauvre ami, je ne vous savais pas si simple!
Comme l'on se trompe pourtant sur la mine! Sous votre masque
froid et méchant, sous vos paroles mordantes, sous la satire
perpétuelle de votre vilain rire, se découvre tout à coup la naïveté
d'un poétereau romantique. Émile, continua-t-elle en s'adressant à
un jeune lycéen qui, la prunelle à la fois allumée et railleuse, suivait
avec intérêt cette conversation, Émile, voulez-vous voir un gobeur?
Regardez monsieur. Ce grand sceptique qui vous paraît peut-être si
fort et si digne de vous servir d'exemple n'est pas autre chose qu'un
gobeur.
Émile fit un geste qui indiquait suffisamment qu'il avait jugé
Réderic.

É
 —Vous ne connaissez pas Émile? poursuivit Julienne. Un petit
cousin à moi, un garçon étonnant. A quinze ans, il vous a des
aperçus stupéfiants sur la vie. Ainsi, tenez, l'autre jour, nous jouions
aux petits papiers. La question posée était celle-ci: «Quelle est la
différence de l'homme et de la femme?» Savez-vous quelle fut la
réponse d'Émile? La voici textuellement: «La différence de l'homme
et de la femme, c'est que la femme descend du singe, tandis que
l'homme y remonte.»
—Est-il possible! se récrièrent les dames avec des gloussements
de rires. Si jeune! Où a-t-il appris ces mots-là? Il n'y a plus
d'enfants!
Le lycéen jouissait avec modestie de son triomphe.
—Voyons, Émile, fit Julienne, puisque vous êtes si précoce,
donnez-nous votre opinion sur M. de Rocrange et Mme Facial.
Émile répondit avec commisération:
—Ils ne sont l'un et l'autre que des serins.
—Un peu osé, pour son âge, mais délicieux! bêla la baronne.
Julienne s'amusait comme une folle.
Sur ces entrefaites, Sénéchal arriva. Il eut un succès d'entrée.
Ces dames l'entourèrent, l'accablèrent de questions.
Une fois assis et les attentions suspendues à ses lèvres:
—Ah! mesdames, débuta-t-il, je sors de l'audience. Quel triste
dénouement! Se peut-il qu'une femme ait pu se résoudre à laisser
traîner devant un tribunal, devant le public, le scandale de sa vie
privée! C'est fait: madame... cette dame... cette femme... je ne sais
plus de quel nom l'appeler... Bref le divorce a été prononcé.
—Contre elle? demanda Réderic.
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