Table Of ContentR. Enay
Palaeontology of
Invertebrates
Translated from the French by Dr. Thomas Reimer
Foreword by Jean Aubouin
With 133 Figures
Springer-Verlag
Berlin Heidelberg New York
London Paris Tokyo
Hong Kong Barcelona
Budapest
Prof. Dr. RAYMOND ENAY
Universite Claude Bernard-Lyon 1
27-43, Bd. du 11 Novembre 1918
F-69622 Villeurbanne Cedex
France
Translated from the French by
Dr. THOMAS REIMER
Via Lavizari 2aIBSM,
CH-6900 Lugano
Switzerland
Title of the Original French edition:
Paleontologie des Invertebres par Raymond Enay, 1990,
©
Bordas, Paris, 1990
ISBN-13:978-3-540-53891-2 e-ISBN-13:978-3-642-76548-3
DOl: 10.1007/978-3-642-76548-3
Library of Congress Cataloging-in-Publication Data. Enay, R. (Raymond) [Paleontologie des
invertebres. English] Palaeontology of invertebrates I R. Enay; translated from the French by
Thomas Reimer; foreword by Jean Aubouin. p. cm. Includes bibliographical references and
index. ISBN-13:978-3-540-5389J-2 88.00F 1. Inverte-
brates, Fossil. I. Title. QE770.E5713 1993 562 - dc20 93-26800
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Foreword
Any explanation of evolution which is oblivious of the
palaeontological information available, will be nothing
more than just a theory in which imagination occupies
a major position.
(P.P. Grasse 1973)
Invertebrate palaeontology has formed the basis of palaeontology
as a whole and, to some extent, also of geology itself. It has led
to the concept of a world that is older than, and different from,
the one in which we are now living.
Although in ancient history fossil shells were of little interest,
the Greek poet Xenophanes of Colophon, who lived during the
sixth century B.C., was the first to record geological observations.
These were taken up and developed further by other classical
scientists and philosophers, and then by Arab scientists like Ibn
Sina (Avicenna) who arrived at a particularly clear view of the
origin of marine sediments.
In contrast to this, in the West it was not until the Renaissance
that the concept of fossils being nothing more than "freaks of
nature" was abandoned and that the most courageous proponents
dared to advocate them as remnants of earlier living organisms.
In this context it was admitted that here and there shells were
found which were similar to extant marine forms and were indica
tive of a different distribution of the seas. This, in turn, cast
doubt on the unchanged existence of the world in its present
state. Among the best intellects willing to consider such releva
tions and starting to doubt the old convictions, albeit only around
the middle of the age of Enlightenment, was Voltaire.
Invertebrate palaeontology was thus in the very centre of
geology right from the start. It led to a palaeogeography which
was quite different from the present reconstructions and prepared
the way for plate tectonics which it adopted without difficulty.
Its increasingly accurate environmental analyses led to palaeo
ecology. As the basis of biostratigraphy, it marks the procession
of time by unraveling a sequence of faunas and floras. It par
ticipates in the struggles on the concept of evolution and on
VI Foreword
human palaeontology itself, an issue which still unleashes fierce
passions. At an early stage it facilitated a quantitative approach
to evolution through the frequently numerous invertebrate fossils
which constitute true popUlations and may be studied as such.
As one of the oldest branches of the earth sciences, together
with mineralogy, it remains a modern science indispensable for
the study of sedimentary sequences.
Raymond Enay, Director of the Centre of Stratigraphic
Palaeontology and Palaeoecology at the University Claude
Bernard at Lyon in France, is eminently suited to compile a
treatise on this subject because of his studies of Mesozoic,
especially Jurassic, biostratigraphy and his numerous reconstruc
tions of the palaeogeographical position of the continents in
the global tectonic framework. In the short and concise book
presented here, both students and informed readers will discover
the richness of the faunas inhabiting the world during earlier
times, and succession of which lends to our modern world the
nature of a mere moment between the past and the future.
JEAN AUBOUIN
Preface
This book, together with Vertebrates Palaeontology by J. Chaline,
forms the second volume of a description of the development of
life on Earth. The volume Micropalaeontology by G. Bignot
describes unicellular animals and plants as well as certain
metazoans of microscopic size. This volume is laid out differently
from the accompanying volume on the vertebrates, not just
for the sake of being different, but because the constraints are
different.
The term invertebrate no longer enjoys the significance of a
group (taxon) of higher rank in the hierarchy of the fossil or
living animal world. It is only a term of convenience to describe
a heterogeneous group which also includes unicellular animals
(protozoans) or is restricted to the metazoans, but it excludes the
vertebrates which are part of the phylum Chordata, a group
which also includes "invertebrate" forms. Despite these limita
tions, the term still covers a wealth of forms. To present such a
vast diversity of forms in a volume comparable in detail to that
on the vertebrates would be beyond the scope of this book and
thus well-nigh impossible.
No less than 20-30 body plans (or organizational levels) are
now known and several have differentiated at an early stage into
distinct structural types. Many are only poorly known or are
unknown from the fossil record, a fact that "luckily" reduces the
number of phyla discussed in this volume to a dozen. After a
short discussion on the origin of life on Earth (Chapter 1) and the
origin of the major body plans (Chapter 2), the remaining phyla
are presented in order of increasing complexity by the most
significant fossil representatives (Chapters 3-8). In each case, the
adaptive significance of the respective forms and/or structures
which more or less profoundly changed the original body plan,
has been treated in detail.
As there are numerous modern treatises which cover
prominently general or thematic aspects of palaeontology, only
one chapter (Chapter 9) is devoted to such aspects and the usual
chapters which appear in the classical treatises, like fossils and
fossilization, the species concept, or palaeontological methods
VIII Preface
and techniques, have been deliberately left out. They are actually
dealt with in the volume on vertebrates, thereby partly establish
ing an equilibrium in the separation of the vertebrates and
invertebrates.
Prof. J. Aubouin, a member of the Institute of France, has
entrusted me with the preparation of this volume for the Geo
sciences Collection for which he is the scientific advisor. I would
like to express my gratitude for his confidence in my abilities and
for the patience he has shown. I would also like to thank Bruno
David, Fran<;oise Debrenne, Christian Gaillard, Jean-Claude
Gall, Jean-Louis Henry, Bernard Lathuiliere and Jean Philip for
their helpful discussions and for providing references and docu
ments throughout the months of preparation of this volume.
Andre Duivon and Arlette Armand prepared the drawings while
typing and correction of the text was carried out by Eleonor
DeOliveira with additional assistance from Noel Podevigne. To
them I would like to extend my sincere gratitude.
Villeurbanne, France RAYMOND ENAY
July 1993
Contents
Chapter 1
The Start of Life on Earth and the First Fossils 1
1.1 Some Introductory Remarks About the Biochemistry
of Living Matter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.1.1 Abilities and Essential Attributes of the
Living Cell. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.1.2 The Fundamental Materials of Vital Processes ....... 2
1.1.3 The Metabolic Processes of Energy Supply . . . . . . . . . . 3
1.1.4 Sources of Energy: Autotrophy and
Heterotrophy .................................... 4
1.2 Physicochemical Model of the Origin of Life. . . . . . . . . 5
1.2.1 The Extraterrestrial Derivation of Life. . . . . . . . . . . . . . 5
1.2.2 The Physicochemical Environment of Terrestrial
Biogenesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
1.2.3 The First Syntheses of Organic Monomers. . . . . . . . . . . 6
1.2.4 The Abiotic Formation of Polymers ................ 7
1.2.5 Cell-Like Assemblages: Microspheres and
Coacervates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
1.3 The Development of Life During the
Precambrian .................................... 10
1.3.1 Evolution Towards an Oxygenic Atmosphere. . . . . . .. 10
1.3.2 Chemical Fossils and Molecular Palaeontology. . . . . .. 13
1.3.3 The First Fossils ................ , . . . . . . . . . . . . . . .. 13
1.4 The Ediacarian or the First Diversified Metazoan
Faunas. . . . . . . . .. . . . . .. . . ... . . .. . . .. . ... . . .. . . .. 15
1.4.1 The Ediacarian Fauna . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15
1.4.2 The Ediacarian Concept and Its Evolutionary
Significance .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17
1.4.3 Palaeoecological Significance . . . . . . . . . . . . . . . . . . . . .. 18
1.4.4 The Later Development of the Ediacarian
Fauna .................................... ,..... 19
Chapter 2
The Invertebrates: Origin and Major Body Plans ....... , . . . 21
2.1 Definition. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 21
x Contents
2.2 Origin and Early Differentiation of the Metazoa 23
2.3 Body Plans of the Lower Invertebrates. . . . . . . . . . . . .. 25
2.3.1 Differentiation of Spongiaria and Coelenterata. . . . . .. 25
2.3.2 Radiation of the Acoelomates ..................... 25
2.4 Radiation of the Coelomates and the Body Plans
of the Higher Metazoans. . . . . . . . . . . . . . . . . . . . . . . . .. 26
2.5 Phanerozoic Evolution and Diversification. . . . . . . . . .. 29
2.5.1 General Faunal Diversification. . . . . . . . . . . . . . . . . . . .. 29
2.5.2 Diversification Within the Phyla ................... 30
Chapter 3
Diploblastic Metazoans: Porifera and Archaeocyatheans . . . .. 33
3.1 Sponges or Porifera. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 33
3.1.1 General Organization ............................ 33
3.1.2 Skeletal Elements and Structures. . . . . . . . . . . . . . . . . .. 35
3.1.3 Fossil Sponges. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 37
3.1.4 Distribution and Habitats. . . . . . . . . . . . . . . . . . . . . . . .. 44
3.2 Archaeocyatheans. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 45
3.2.1 Organization and Structure. . . . . . . . . . . . . . . . . . . . . . .. 45
3.2.2 Classification and Evolutionary Trends. . . . . . . . . . . . .. 49
3.2.3 Distribution and Habitats ........................ ~ 50
Chapter 4
Diploblastic Metazoans: The Coelenterata. . . . . . . . . . . . . . . .. 52
4.1 Major Body Plans and Origin of the Cnidaria . . . . . . .. 52
4.1.1 Basic Structural Plan. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 52
4.1.2 The Three Classes of Cnidaria . . . . . . . . . . . . . . . . . . . .. 53
4.1.3 Origin and Affiliations of the Cnidaria . . . . . . . . . . . . .. 53
4.1.4 Adaptive Radiation of the Cnidaria. . . . . . . . . . . . . . . .. 55
4.2 Hydrozoans and Scyphozoans ..................... 56
4.2.1 The Hydrozoans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 56
4.2.2 Fossil Scyphozoans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 57
4.3 The Anthozoans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 59
4.3.1 The Anthozoan Body Plan....... . . . ....... . . ..... 60
4.3.2 The Cerianthipatharia . . . . . . . . . . . . . . . . . . . . . . . . . . .. 60
4.3.3 The Octocorallia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 60
4.3.4 The Zoantharia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 62
Chapter 5
Triploblastic Pseudometamerous Coelomates: The Molluscs.. 73
5.1 Origin and Diversification of the Different Classes
of Molluscs ..................................... 73
Contents XI
5.1.1 The Mollusc Type or Protomollusc . . . . . . . . . . . . . . . .. 73
5.1.2 Initial Early Radiation. . . . . . . . . . . . . . . . . . . . . . . . . . .. 74
5.1.3 The Oldest Fossil Molluscs. . . . . . . . . . . . . . . . . . . . . . .. 76
5.2 Polyplacophorans (Loricata) and
Monoplacophorans ............................... 76
5.2.1 The Polyplacophorans (Loricata). . . . . . . . . . . . . . . . . .. 77
5.2.2 The Monoplacophorans . . . . . . . . . . . . . . . . . . . . . . . . . .. 77
5.3 The Gastropods ................................. 78
5.3.1 The Basic Organization of a Gastropod . . . . . . . . . . . .. 78
5.3.2 The Gastropod Shell. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 80
5.3.3 Palaeontological History of the Gastropods. . . . . . . . .. 82
5.3.4 Heteropods and Pteropods. . . . . . . . . . . . . . . . . . . . . . .. 85
5.4 Rostroconchs and Scaphopods . . . . . . . . . . . . . . . . . . . .. 85
5.4.1 The Rostroconchs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 85
5.4.2 The Scaphopods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 86
5.5 Bivalves (or Lamellibranchs, Pelecypods,
Acephalia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 87
5.5.1 Typical Body Plan of the Bivalves . . . . . . . . . . . . . . . . .. 87
5.5.2 Characteristic Aspects of the Bivalve Shell:
Hinge and Ligament. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 90
5.5.3 Functional Morphology of the Shell ................ 95
5.5.4 Adaptive Radiation of the Bivalves ................. 100
5.6 Cephalopods .................................... 104
5.6.1 The Major Cephalopod Groups and Their
Relationships. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 105
5.6.2 The Nautiloids .................................. 108
5.6.3 The Bactritoids and the Origin of the
Ammonoids. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 116
5.6.4 The Ammonoids ................................. 117
5.6.5 Phylogenetic History of the Ectocochlian
Cephalopods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 124
5.6.6 The Coleoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 131
Chapter 6
Triploblastic Eumetamerous Coelomates: The Arthropods . .. 136
6.1 Major Distinctive Features of the Arthropods. . . . . . .. 137
6.1.1 The First Fossil Arthropods ....................... 137
6.1.2 The Carapace and the Arthropod Segments ......... 138
6.1.3 The Paired Appendages of the Arthropods . . . . . . . . .. 139
6.1.4 The Major Body Sections of the Arthropods ......... 140
6.1.5 The Arthropods: A Monophyletic Group or a
Functional Degree of Organization? ................ 142
6.2 Arthropod Radiation in the Aquatic Environment:
Trilobites and Crustaceans ........................ 145
